Research Article |
Corresponding author: Daniel Juřena ( aphodius@seznam.cz ) Academic editor: Andrey Frolov
© 2022 Daniel Juřena.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Juřena D (2022) A critical review of the distribution of the endangered European earth-borer beetle Bolbelasmus unicornis (Coleoptera, Geotrupidae), with new records from 13 countries and observations on its bionomy. ZooKeys 1105: 1-125. https://doi.org/10.3897/zookeys.1105.81474
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The distribution of Bolbelasmus unicornis (Schrank, 1789) is critically reviewed throughout its range with emphasis on the Czech Republic and Slovakia. The species has been reliably recorded from 377 localities in 19 countries. New records are given from 152 localities of Bulgaria, Czech Republic, Germany, Hungary, Italy, Moldova, Poland, Romania, Serbia, Slovakia, Turkey, and Ukraine. For Germany, the species is recorded for the first time in 54 years. The occurrence of the species in Switzerland is confirmed by two historical specimens from Zürich. The only known historical specimen labelled “Kaukasus” is given, which could originate from Russia, where this species has not been recorded before (however, confusion of the locality label cannot be ruled out). All published faunistic data from across the range are presented here in full, in several cases supplemented by details subsequently obtained by the author. Distribution maps are compiled separately for the Czech Republic and Slovakia, and for the entire range. A separate map is also available for Hungary, where approximately one-third of the known localities are located. Statistical data concerning the flight activity of adults, seasonal dynamics for part of the distribution area, details of records and notes on the bionomy and ethology of the species are provided. Possible feeding strategies for adults and larvae of B. unicornis are discussed, as well as current knowledge of the natural history of various representatives of the subfamily Bolboceratinae. A monitoring method for the species is proposed.
Asia Minor, Bolboceratinae, ethology, Europe, Palaearctic realm, zoogeography
Bolbelasmus unicornis (Schrank, 1789) is a European species of earth-borer beetle extending into the western Asian part of Turkey with the centre of distribution in the Pannonian Basin (see Faunistic records and Fig.
The species was described as Scarabaeus unicornu by
Given its secretive lifestyle and lack of knowledge of effective collecting methods, the distribution and bionomy of B. unicornis are poorly known. Adults spend most of their time underground, with above-ground activity limited to short flight periods when they fly very close to the ground just after sunset (see Natural history of Bolboceratinae in this study). Nothing is known about the immature stages and the diet of adults and larvae. However, some authors assumed that both adults and larvae feed on hypogeous fungi (e.g.,
The nomenclature used in this research follows
Faunistic records from the Czech Republic and Slovakia are divided into paragraphs beginning with a number representing the code of the faunistic square that refers to the Central European grid for mapping fauna and flora (Fig.
The following systems are used to transliterate cited literature and geographical or personal names in the Cyrillic and Armenian scripts: BGN/PCGN 2013 Agreement for Bulgarian, BGN/PCGN 1947 System for Russian, BGN/PCGN 2005 Agreement for Serbian, BGN/PCGN 2019 Agreement for Ukrainian, and BGN/PCGN 1981 System for Armenian.
For the distribution map of the Czech Republic and Slovakia, the records are divided into three time periods: the records before 1960, records between 1960–1999, and records after 1999 (Fig.
Statistics on flights of adults were compiled for eight localities (seven Slovak and one Serbian), for which detailed data were available (Tables
The graph of seasonal dynamics was generated with data obtained from countries of the Pannonian Basin for which data on a minimum of 30 specimens were available (Fig.
The dates of Panzer’s works are adopted from
ABC Attila Balázs, Čamovce, Slovakia
ABZ Andrii Ivanovych Bachynskyi (Андрій Іванович Бачинський), Zalishchyky, Ukraine
ADW Alexander Dostal, Vienna, Austria
AGB András Gór, Biatorbágy, Hungary
AHB Adam Hergovits, Bratislava, Slovakia
AKB Attila Kotán, Budapest, Hungary
AMK András Máté, Kecskemét, Hungary
APC Alexandru-Mihai Pintilioaie, Comănești, Romania
APE Attila Pál, Érd, Hungary
APO Antonín Peutlschmid, Olomouc, Czech Republic
ARC Adrian Ruicănescu, Cluj-Napoca, Romania
ASH Aleš Sedláček, Hranice, Czech Republic
ASK Artur Anatoliiovych Shekhovtsov (Артур Анатолійович Шеховцов), Kharkiv, Ukraine
AUP Ákos Uherkovich, Pécs, Hungary
BBO Boris Bubeník Sr., Ostrava, Czech Republic
BCK Csaba Bán, Kecskemét, Hungary
BJN Jiří Brestovanský Jr., Neratovice, Czech Republic
BJO Boris Bubeník Jr., Ostrava, Czech Republic
BKL Bence Krajcsovszky, Lábatlan, Hungary
BMP Marek Bunalski, Poznań, Poland
BSP † Svatopluk Bílý, Prague, Czech Republic
BVK Bohdan Mykolaiovych Vasko (Богдан Миколайович Васько), Kyiv, Ukraine
CBE Csaba Bartha, Eger, Hungary
CBK Csaba Bíró, Kecskemét, Hungary
CKZ Csaba Kutasi, Zirc, Hungary
CMI Cosmin-Ovidiu Manci, Iași, Romania
CSB Csaba Szabóky, Budapest, Hungary
CSS Csaba Szinetár, Szombathely, Hungary
CVK Csaba Vadász, Kecskemét, Hungary
CWP Christian Wieser, Pischeldorf, Austria
DCO Dan Čagánek, Otrokovice, Czech Republic
DHH David Hrebeň, Havířov, Czech Republic
DHP David Horák, Prostějov, Czech Republic
DJP Daniel Juřena, Prostějov, Czech Republic
DKC Denis Keith, Chartres, France
DKP David Král, Prague, Czech Republic
DPB Dragan Pavićević (Драган Павићевић), Belgrade, Serbia
DPS Dmytro Protopopov (Дмитро Протопопов), Semyhiria, Ukraine
DRW Dominik Rabl, Vienna, Austria
DVB Dalibor Všianský, Brno, Czech Republic
DVH Dušan Vacík, Hranice, Czech Republic
DVZ Daniel Vít, Zlín, Czech Republic
EJB Eduard Jendek, Bratislava, Slovakia
FKD Ferenc Klecska, Dunaharaszti, Hungary
FPT Filip Pavel, Týniště nad Orlicí, Czech Republic
FSB Filip Štrba, Bratislava, Slovakia
FSP František Štěpánek, Přerov, Czech Republic
FTK Florian Theves, Weingarten (Baden), Germany
FTR Filip Trnka, Rychnov nad Kněžnou, Czech Republic
FTV Filip Trojan, Velké Němčice, Czech Republic
GAB Ádám Gór, Biatorbágy, Hungary
GPB Gergely Petrányi, Budapest, Hungary
GML Geoffrey Miessen, Liège, Belgium
GSB Győző Szél, Budapest, Hungary
HDO Hryhorii Mykolaiovych Demydov (Григорій Миколайович Демидов), Odessa, Ukraine
HMS Heinz Mitter, Steyr, Austria
HTR Hennadii Tarasenko (Геннадій Тарасенко), Rzhyshchiv, Ukraine
IIB Ionuț Ștefan Iorgu, Bucharest, Romania
IJN Ivo Jeniš, Náklo, Czech Republic
IMO Ivo Martinů, Olomouc, Czech Republic
IMP Ivan Marvan, Pardubice, Czech Republic
IPO Ivan Paloušek, Olomouc, Czech Republic
IRB Imre Retezár, Budapest, Hungary
ITV Ilja Trojan, Velké Němčice, Czech Republic
JAH † Josef Adámek, Hradec Králové, Czech Republic
JBB Jan Bezděk, Brno, Czech Republic
JCM Josef Chybík, Modřice, Czech Republic
JDB János Dobos, Budapest, Hungary
JDC Jiří Dvořák, Čelčice, Czech Republic
JHH Jiří Hájek, Hrobce, Czech Republic
JHL Jan Helešic, Lužice (near Hodonín), Czech Republic
JHM Jaromír Hanuš, Moravské Budějovice, Czech Republic
JHP Jiří Hanzlík, Přerov–Popovice, Czech Republic
JJP † Josef Jurčíček, Prague, Czech Republic
JKB Ján Kodada, Bratislava, Slovakia
JKJ Jaroslav Kaláb, Jinačovice, Czech Republic
JKO Jindřich Kuja, Olomouc, Czech Republic
JKP Josef Krošlák, Plzeň, Czech Republic
JKV Josef Kadlec, Varnsdorf, Czech Republic
JMB József Muskovits, Budapest, Hungary
JMD † Jaroslav Máslo, Dolní Radechová, Czech Republic
JMH Jan Matějíček, Hradec Králové, Czech Republic
JPH Jan Pelikán, Hradec Králové, Czech Republic
JPP Jiří Plecháč, Pecka, Czech Republic
JRC Jindřich Ryšavý Jun., České Budějovice, Czech Republic
JRS Jaroslav Resl, Sněžné (near Nové Město nad Metují), Czech Republic
JSP Jan Schneider, Prague, Czech Republic
JST József Sár, Teklafalu, Hungary
JSU Jiří Spružina, Ústí nad Labem, Czech Republic
JTK Jaroslav Turna, Kostelec na Hané, Czech Republic
JVP † Jan Vitner, Prague, Czech Republic
JZJ Jaroslav Žák, Jezernice, Czech Republic
KBB Karol Bucsek, Bratislava, Slovakia
KDO Karel Doležel, Olomouc, Czech Republic
KFW Katrin Fuchs, Vienna, Austria
KHE Krisztián Harmos, Eger, Hungary
KLP Jiří Klícha, Prague, Czech Republic
KPH Karel Pils, Hlohovec, Slovakia
KPL Krzysztof Pałka, Lublin, Poland
KPV † Karel Poláček, Vysoké Mýto, Czech Republic
KRU Květoslav Resl, Uherský Brod, Czech Republic
KVB Klára Varga Bánné, Kecskemét, Hungary
KVM Kirill Vladimirovich Makarov (Кирилл Владимирович Макаров), Moscow, Russia
KVS Václav Křivan, Štěměchy, Czech Republic
LAB László Ádám, Budapest, Hungary
LEN Ladislav Ernest, Nymburk, Czech Republic
LFI Lucian Fusu, Iași, Romania
LHI Lucian Hănceanu, Iași, Romania
LKK Ladislav Kandrnál, Kunovice (near Uherské Hradiště), Czech Republic
LKM Ľudevít Kašovský, Martin, Slovakia
LMB Levente Erik Moharos, Budakeszi, Hungary
LMN Ladislav Miško, Nové Zámky, Slovakia
LMO Luboš Mazal, Olomouc, Czech Republic
LMT Ladislav Mencl, Týnec nad Labem, Czech Republic
LNB László Nádai, Budapest, Hungary
LRL Luciano Ragozzino Lerma, Italy
MBO Michal Bednařík, Olomouc, Czech Republic
MBF Marco Bastianini, Follonica, Italy
MBP Milan Brabec, Prague, Czech Republic
MHP Marcel Hájek, Plzeň, Czech Republic
MJR Martin Jagelka, Rohožník, Slovakia
MKJ Martin Kejzlar, Jevíčko, Czech Republic
MKY Mark Yuriyevich’ K’alashyan (Մարկ Յուրիևիչ Քալաշյան), Yerevan, Armenia
MLB Márk Lukátsi, Budapest, Hungary
MLS Miloš Lošťák, Šumperk, Czech Republic
MNB Martin Němec, Brno, Czech Republic
MNR Milan Nikodým, Roztoky (near Prague), Czech Republic
MOB Oto Majzlan, Bratislava, Slovakia
MPK Michal Pikner, Kněžpole (near Uherské Hradiště), Czech Republic
MPN Miloš Popović (Милош Поповић), Niš, Serbia
MPP Milan Sláma, Prague, Czech Republic
MPV Milan Pilarčík, Velké Pavlovice, Czech Republic
MRM Miklós Ringler, Munich, Germany
MRV Milan Ryšavý, Vlkoš (near Přerov), Czech Republic
MSB Milan Štrba, Bratislava, Slovakia
MSN Marko Šćiban (Марко Шћибан), Novi Sad, Serbia
MSZ Miloslav Šanda, Žatec, Czech Republic
MTM Maximilian Teodorescu, Măgurele, Romania
MTS Milan Trubačík, Staré Město (near Uherské Hradiště), Czech Republic
MVP Martin Volf, Prague, Czech Republic
MYP Vladislav Malý, Prague, Czech Republic
MZB Miroslav Zúber, Bradlec, Czech Republic
MZK Miroslav Ihorovych Zaika (Мирослав Ігорович Заїка), Kyiv, Ukraine
MZP Zdena Martinová & Zdeněk Znamenáček, Prague, Czech Republic
NKB Naďa Kazdová, Brno, Czech Republic
NPB Norbert Pometkó, Budapest, Hungary
OHS Oliver Hillert, Schöneiche bei Berlin, Germany
OKO Oleksandr Oleksandrovych Kolomeichuk (Олександр Олександрович Коломейчук), Odessa, Ukraine
OMB † Ottó Merkl, Budapest, Hungary
OSD Oleksandr Oleksiiovych Sukhenko (Олександр Олексійович Сухенко), Dnipro, Ukraine
OSO Ondřej Sabol, Ostrava–Nová Bělá, Czech Republic
OVK Oleksii Vasyliuk (Олексій Василюк), Kyiv, Ukraine
PCB Petr Čechovský, Brno, Czech Republic
PFS Péter Farkas, Ságvár, Hungary
PIL Pavel Imríšek, Louny, Czech Republic
PJH Josef Pavlas, Havířov, Czech Republic
PJL Pavel Jáchymek, Luhačovice, Czech Republic
PKB Péter Kovács, Budapest, Hungary
PKG Peter Kurina, Gajary, Slovakia
PMB Peter Mihálik, Bratislava, Slovakia
PPB Petr Pacholátko, Brno, Czech Republic
PSZ Petr Středulínský, Zlín, Czech Republic
PVP Petr Včelička, Prague, Czech Republic
PZW Petr Zábranský, Vienna, Austria
RCP † Radek Červenka, Prague, Czech Republic
RCR Roman Cséfalvay, Rohovce–Kyselica, Slovakia
REE Róbert Enyedi, Eger, Hungary
RFO † Rostislav Fornůsek, Olomouc, Czech Republic
RGM Rudolf Gabzdil, Michalovce, Slovakia
RHB Roman Hergovits, Bratislava, Slovakia
RHK Rostyslav Pavlovych Herasymov (Ростислав Павлович Герасимов), Kyiv, Ukraine
RKP Radim Klíč, Prostějov, Czech Republic
RMU Robert Machálek, Uherské Hradiště, Czech Republic
RPM Riccardo Pittino, Milan, Italy
RSG Roland Štefanovič, Galanta, Slovakia
RSP Radim Šigut, Paskov, Czech Republic
RVO † Radovan Veigler, Olomouc, Czech Republic
RZJ Renata Žáková, Jezernice, Czech Republic
SBP Sándor Bérces, Pomáz, Hungary
SBS Stoyan Beshkov, Sofia, Bulgaria
SDP Stanislav Doležal, Plzeň–Božkov, Czech Republic
SIB Sándor Ilniczky, Budapest, Hungary
SKP Jiří Skýpala, Prague, Czech Republic
SPP Serge Peslier, Perpignan, France
SRB Richard Schnürmacher, Bratislava, Slovakia
SRL Stefan Rabl, Lengenfeld, Austria
STK Serhii Vasylovych Tsykal (Сергій Васильович Цикал), Kyiv, Ukraine
SZM Stefano Ziani, Meldola, Italy
TBK Torsten Bittner, Weißenberg, Germany
TDB Tamás Deli, Békéscsaba, Hungary
TDS Tibor Danyik, Szarvas, Hungary
TGK Tomáš Grulich, Dolní Studénky–Králec, Czech Republic
TKB † Tibor Kovács Sr., Bátonyterenye–Kisterenye, Hungary
TKG Tibor Kovács Jr., Gyöngyös, Hungary
TKH Tomáš Kopecký, Hradec Králové, Czech Republic
TKK Tamás Kiss, Kecskemét, Hungary
TNB Tamás Németh, Gödöllő, Hungary
TSH Tibor Spevár, Hlohovec, Slovakia
TVP Tomáš Vendl, Prague, Czech Republic
VDP Václav Dongres, Plzeň, Czech Republic
VGG † Vadim Gennad’yevich Grachëv (Вадим Геннадьевич Грачёв), Moscow, Russia
VJP Vojtěch Jiříček, Prostějov, Czech Republic
VKK Vitalii Volodymyrovych Kavurka (Віталій Володимирович Кавурка), Kyiv, Ukraine
VKS Vítězslav Kubáň, Šlapanice, Czech Republic
VLP Jaromír Vališ, Přerov, Czech Republic
VMP Vlastimil Mihal, Přerov, Czech Republic
VPB Vilmos Polonyi, Budapest, Hungary
VRH Vladislav Řebíček, Hradištko, Czech Republic
VSI Valentin Szénási, Isaszeg, Hungary
VSK Vasyl Musiiovych Serhiienko (Василь Мусійович Сергієнко), Kyiv, Ukraine
VSM Viktor Viktorovych Strenada (Віктор Вікторович Стренада), Mykolaiv, Ukraine
VTH Vladimír Thomka, Humenné, Slovakia
VTO Viacheslav Anatoliiovych Trach (Вячеслав Анатолійович Трач), Odessa, Ukraine
VUB Viorel Ungureanu, Buzău, Romania
VVO Vladimír Vyhnálek, Olomouc, Czech Republic
VZO Vladimír Zeman, Olomouc, Czech Republic
WBW Wolfgang Barries, Vienna, Austria
WHS Walter Heinz, Schwanfeld, Germany
YHS Yurii Mykolaiovych Geriak (Юрій Миколайович Ґеряк), Sambir, Ukraine
YKL Yurii Vasylovych Kanarskyi (Юрій Васильович Канарський), Lviv, Ukraine
YKO Yevhenii Vitaliiovych Khalaim (Євгеній Віталійович Халаїм), Odessa, Ukraine
YSK Yurii Yevhenovych Skrylnyk (Юрій Євгенович Скрильник), Kharkiv, Ukraine
ZBB Zoran Božović (Зоран Божовић), Batajnica, Serbia
ZBP Zsófia Mocskonyi Bércesné, Pomáz, Hungary
ZCP Zdeněk Čermák, Prostějov, Czech Republic
ZDP † Zdeněk Doležal, Plzeň, Czech Republic
ZKB Zoltán Körmendy, Budapest, Hungary
ZKM Zdeněk Kraus, Mikulovice (near Znojmo), Czech Republic
ZLB Zdeněk Laštůvka, Brno, Czech Republic
ZVP Zdeněk Vancl, Police nad Metují, Czech Republic
BNMS Brukenthal National Museum, Sibiu, Romania
BZLA Biologiezentrum Linz, Austria
CUIR Alexandru Ioan Cuza University, Iași, Romania
ETHZ Entomological collection of the Swiss Federal Institute of Technology, Zürich, Switzerland
FGBI Franziskaner Gymnasium Bozen, Bolzano, Italy
GANM “Grigore Antipa” National Museum of Natural History, Bucharest, Romania
GUNU Nizhyn Gogol State University, Nizhyn, Ukraine
JHIS Jovan Hadži Institute of Biology of the Research Centre of the Slovenian Academy of Sciences and Arts, Ig, Slovenia
LKKA Landesmuseums für Kärnten, Klagenfurt am Wörthersee, Austria
MHKC Museum of Eastern Bohemia in Hradec Králové, Hradec Králové, Czech Republic
MJMC Muzeum jihovýchodní Moravy ve Zlíně, Zlín, Czech Republic
MKPC Muzeum Komenského v Přerově, Přerov, Czech Republic
MNBG Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Museum für Naturkunde, Berlin, Germany
MMGH Mátra Museum of the Hungarian Natural History Museum, Gyöngyös, Hungary
MNFI Natural History Museum “La Specola”, Florence, Italy
MNSA Museum Niederösterreich, Sankt Pölten, Austria
MPGU Moscow Pedagogical State University, Moscow, Russia
MSNB Museo di Scienze Naturali dell’Alto Adige, Bolzano, Italy
MTDG Senckenberg Naturhistorische Sammlungen, Museum für Tierkunde, Dresden, Germany
NHMK State Natural History Museum of V. N. Karazin Kharkiv National University, Kharkiv, Ukraine
NHMU National Science and Natural History Museum of the National Academy of Sciences of Ukraine, Kyiv, Ukraine
NMCM National Museum of Ethnography and Natural History, Chișinău, Republic of Moldova
RMNH Naturalis Biodiversity Centre (formerly Rijksmuseum van Natuurlijke Historie), Leiden, Netherlands
SMLU State Museum of Natural History, Lviv, Ukraine
SNMS Slovak National Museum–Natural History Museum, Bratislava, Slovakia
TMLS Tekovské múzeum v Leviciach, Levice, Slovakia
VMHS Vihorlatské múzeum Humenné, Slovakia
ZMNU Zoological Museum of the Taras Shevchenko National University, Kyiv, Ukraine
ZMPC Západočeské muzeum v Plzni, Plzeň, Czech Republic
ZINR Zoological Institute of Russian Academy of Sciences, Saint Petersburg, Russia
ZUDH Department of Nature Conservation, Zoology and Game Management, University of Debrecen, Debrecen, Hungary
Family: GEOTRUPIDAE Latreille, 1802
Subfamily: Bolboceratinae Mulsant, 1842
Tribe: BOLBELASMINI Iablokoff-Khnzorian, 1977
Genus: Bolbelasmus Boucomont, 1911
Subgenus: Bolbelasmus Boucomont, 1911
Species: Bolbelasmus (Bolbelasmus) unicornis (Schrank, 1789)
Published data
? 5354: “Kummer” [= Hradčany near Mimoň], 1 ♂ flying in the evening, no other data (
5756: Loučeň, 28.v.1905, 1 spec., [Augustin] Šrámek leg., Radek Červenka and Radek Dunda det., coll.
6865: “Kammberg b. Brünn” [probably Brno – Kohoutovice env., perhaps Kamenný vrch hill], no other data (
7067: Hovorany, 6.v.1941, Jan Roubal leg. (
6568: Prostějov, [between 1878–1899, see
? 6570: Přerov env. [probably Bochoř near Přerov], no other data (
6870: “Ungarisch Hradisch” [= Uherské Hradiště] env., Morava River valley, no other data (
Material examined and new observations
7067: Bořetice env., PR Zázmoníky, 48°56'06.9"N, 16°51'20.5"E, ca. 300 m a.s.l., 1.v.1998, 1 elytron excavated from loess soil, KRU obs.; Čejč env., “Květnatá step” [= so-called Květnatá steppe, northern part of the PR Čejkovické Špidláky reserve], 48°55'22.0"N, 16°57'24.2"E, ca. 190 m a.s.l., 1.vii.1995, remains of a female excavated from a burrow of Oryctolagus cuniculus, KRU (VKS det., 15 October 2005); Čejč env., “Mansonova step” [= so-called “Manson’s steppe”], 48°55'32.1"N, 16°58'46.6"E, ca. 210 m a.s.l., 20.vi.1986, 1 ♀ FSLG after sunset, PCB; 17.vi.1988, 1 ♀ FSLG after sunset, PCB; 21.vi.1988, 2 ♀♀ FSLG after sunset, VKS; 27.vi.1988, 8 spec. FSLG after sunset (for a photograph of one of them see
6376: “Friedek Umg.” [= Frýdek-Místek env.], 20.vi.1923, 1 ♀ [ex. coll. Dr Karel Samšiňák], Jos[ef] Hlisnikowski [leg.], DJP det., coll.
Comment
In the Czech Republic, the species is known from a few localities only. Old reports by
Published data
7867: [Bratislava env.,] Děvín: Kobyla [= Devínska Kobyla hill], [between 1921–1936, see
7568: Malacky, no other data (
7868: “Pozsony” or “Presburg” [= Bratislava] env., no other data (
7968: Bratislava, Kopáč Island, [PR Kopáčsky ostrov], 19.v.2006, 1 spec., Malaise trap, MOB leg. (
7969: “Somorja” [= Šamorín], 10.v.1897, 1 spec. inside the digestive system of Upupa epops, Ernő Csiki obs. (
7371: “Pustá Ves” [= Prašník – Horná Pustá Ves or Dolná Pustá Ves], 22.vii.1984, 1 ♀, [at light], JMD leg., coll.
7272: Čachtice, [probably between 1920–1938, see
7572: Hlohovec, undated [probably first half of the 20th century], 1 ♀, Várkonyi leg., coll. DKP deposited in
7373: Brunovce, no other data, 1 ♀ in coll.
7074: “Liborcsa” [= Nemšová – Ľuborča], [ca. 230 m a.s.l.], undated [probably second half of the 19th century], 2 spec., Nitnausz leg. (
7174: Trenčín, date not specified, old vineyard, more spec, Rudolf Čepelák, František Hajný, and Ladislav Korbel leg., and 1 spec., Jan Roubal leg. (
7274: “Trenčín – Inovec” [= Považský Inovec Mts, Inovec hill env.], undated [probably 1920s-1930s], 1 ♂, [Rudolf] Čepelák leg., ex original coll. Rudolf Veselý, currently in coll.
7374: “Podhragy” [= Podhradie near Topolčany], June and July 1895–1897, collector not specified (
8174: “Keszegfalu” [= Keszegfalva, currently Kameničná], 25.v.1906, 1 spec. inside the digestive system of Falco vespertinus, Ernő Csiki obs. (
7275: Ľutov [env., Pálenice hill, ca. 48°46'57"N, 18°16'44"E, 250–300 m a.s.l.], 1.vi.–15.vii.[probably 1920s], Čepelák leg. (
8176: “Bátorkeszi” [= Bátorove Kosihy], June [between 1919–1923, see
8177: Štúrovo (8278) [Štúrovo env., Belianské kopce hills, Modrý vrch hill env., PR Vŕšok env.], 24.v.1985, 2 spec., RFO (
8178: Kamenica nad Hronom env., 47°50'29.5"N, 18°43'34.8"E, 9.vii.1980, ca. 30 spec. FSLG around midnight, PJH leg., 1 ♂ and 1 ♀ in coll. VJP, 1 ♀ in coll. ZDP deposited in ZMPC (
8179: Chľaba env., 47°49'27"N, 18°50'57"E (the site near the confluence of the Danube and Ipeľ rivers), 103 m a.s.l., 5.vii.1975, plant materials alluviated by flooded Danube and Ipeľ rivers, 1 ♀, VKS leg. et coll., 1 ♂, PPB leg., coll. VKS (
7781: “Plachti[n]ce” [= Horné, Stredné or Dolné Plachtince], 5.vi.1938, [Rudolf] Schwarz leg. (
7683: “Losoncz” [= Lučenec], 1877–1891, Emil Malesevics leg. (
7884: Šiatorská Bukovinka, parking at the cemetery, [48°11'4"N, 19°49'33"E; 290 m a.s.l.], 8.vii.1973, 1 spec., at light (kerosene lamp), SKP leg., coll. SPP (
7785: Hajnáčka [– Buková, 48°13'36.97"N, 19°58'24.11"E, steppe slope near the forest], 15.vii.1984, 1 ♀, dead on the ground, RCP (
7882: Kiarov, 15.–20.vi.1936, 1 ♀, [Dr Rudolf] Schwarz leg., ex original coll. Bohumil Štícha, currently in coll.
7277: Prievidza, forest park, 18.vii.1995, 1 ♂, RGM; (
7280: Banská Bystrica, 18.v.1979, 1 ♂, collector unknown, coll. KVS (
7488: Silická Brezová, 3.vi.1999, 1 ♀, dead on a path crossing a steppe meadow, KDO (
7390: Hrhov, 20.–21.vii.1981, 10 spec. excavated from their burrows, in a few cases together with Od. armiger (at a depth of up to 7 cm, the burrows changed direction from vertical to horizontal; in two cases, in one hole were two males or two females of B. unicornis together), LMT (
7494: Slanská Huta env., 48°34'54.8"N, 21°28'31.7"E, 600 m a.s.l., 24.vii.1972, 1 ♀ crawling on the ground after sunset, ZLB obs. + photo – see Fig.
7596: Ladmovce, 9.viii.1982, 2 ♂♂ excavated from their burrows from a depth of 8 cm, and 1 ♀ from a depth of ca. 20 cm, LMT (
7097: Lackovce env., Veľká hill, [ca. 48°56'35"N, 21°58'13.5"E], 2.vii.–31.viii.2001, 2 ♂♂ and 2 ♀♀, steep forest-steppe hillside with shrubbery of Rosa canina and Prunus spinosa, pitfall traps with formaldehyde, together with more spec. of Od. armiger, VTH leg., coll. VMHS (
7098–7099: Snina, July 1965, 1 ♂, MPP leg., coll. DKP deposited in
Material examined and new observations
7868: “Pressburg” [= Bratislava], no other data, 2 ♂♂ and 3 ♀♀ in coll.
7868–7869: Bratislava – Podunajské Biskupice, Kopáč Island, PP Panský diel env. (Figs
Data on flights of adults of B. unicornis at the locality of PP Panský diel. Key: BF = beginning of flights, EF = end of flights, S = sunset, S-BF = time period from sunset to the beginning of flights, S-EF = time period from sunset to the end of flights, DF = duration of flights, T = air temperature during flights.
Slovakia, Bratislava – Podunajské Biskupice, Kopáč Island, PP Panský diel | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
7.vi.2016 | 11 (6/5) | 21.25 | 21.45 | 20.47 | 38 min | 58 min | 20 min | 14 °C | heavy dew, no wind |
8.vi.2016 | 4 (4/-) | 21.27 | 21.43 | 20.48 | 39 min | 55 min | 16 min | 16 °C | dew, no wind |
18.vi.2016 | 6 (3/3) | 21.33 | 21.53 | 20.54 | 39 min | 59 min | 20 min | 19 °C | dew, no wind |
21.vi.2016 | 5 (3/2) | 21.31 | 22.03 | 20.54 | 37 min | 69 min | 32 min | 21 °C | no wind |
22.vi.2016 | 6 (2/4) | 21.21 | 21.57 | 20.55 | 26 min | 62 min | 36 min | 22 °C | no wind |
23.vi.2016 | 11 (6/5) | 21.33 | 21.53 | 20.55 | 38 min | 58 min | 20 min | 23 °C | no wind |
24.vi.2016 | 24 (10/14) | 21.36 | 22.08 | 20.55 | 41 min | 73 min | 32 min | 25 °C | no wind |
25.vi.2016 | 7 (2/5) | 21.28 | 22.08 | 20.55 | 33 min | 68 min | 32 min | 26 °C | no wind |
26.vi.2016 | 25 (13/12) | 21.31 | 22.09 | 20.55 | 36 min | 74 min | 38 min | 20 °C | light breeze |
27.vi.2016 | 10 (2/8) | 21.31 | 21.57 | 20.55 | 36 min | 62 min | 26 min | 20 °C | gentle breeze |
28.vi.2016 | 9 (2/7) | 21.28 | 21.48 | 20.55 | 33 min | 53 min | 20 min | 19 °C | light breeze |
29.vi.2016 | 14 (8/6) | 21.29 | 22.03 | 20.55 | 34 min | 68 min | 34 min | 24 °C | light dew, no wind |
21.vii.2016 | 36 (25/11) | 21.09 | 21.51 | 20.41 | 28 min | 70 min | 42 min | 24 °C | no wind |
22.vii.2016 | 51 (28/23) | 21.08 | 21.51 | 20.40 | 28 min | 71 min | 43 min | 23 °C | ground mist, no wind |
23.vii.2016 | 41 (26/15) | 21.09 | 21.53 | 20.39 | 30 min | 74 min | 44 min | 24 °C | light ground fog, no wind |
24.vii.2016 | 68 (38/30) | 21.08 | 21.49 | 20.38 | 30 min | 71 min | 41 min | 23 °C | dew, no wind |
25.vii.2016 | 15 (14/1) | 21.06 | 21.41 | 20.37 | 29 min | 64 min | 35 min | 22 °C | light rain, no wind |
29.vii.2016 | 13 (10/3) | 21.02 | 21.30 | 20.32 | 30 min | 58 min | 28 min | 22 °C | light dew, no wind |
30.vii.2016 | 37 (16/21) | 20.53 | 21.56 | 20.30 | 23 min | 86 min | 63 min | 24 °C | light dew, no wind |
7.viii.2016 | 29 (16/13) | 20.46 | 21.16 | 20.19 | 27 min | 57 min | 30 min | 17 °C | no wind |
8.viii.2016 | 46 (26/20) | 20.45 | 21.25 | 20.17 | 28 min | 68 min | 40 min | 16 °C | no wind |
13.viii.2016 | 51 (32/19) | 20.40 | 21.01 | 20.09 | 31 min | 52 min | 21 min | 19 °C | no wind |
n (♂/♀) | 519 (292/227) | average | 32 min | 65 min | 32 min | 21 °C |
Bratislava, Kopáč Island (Slovakia), the area with the largest known population of B. unicornis in Europe before its conversion by inappropriate conservation management (removal of small trees and shrubs and introduction of intensive sheep grazing) A view of the site in 1950 B view of the site in 2014 with the area of the highest density of B. unicornis outlined with yellow borders C detail of the area with the highest density of the species with faunistic squares marked (see Materials and methods and Fig.
7968: Rusovce – Záhrady, 48°3'20.614"N, 17°9'18.014"E, 140 m a.s.l., 6.vi.2020, 1 ♂ flying ca. 10–20 cm above the ground at 21.20 CEST, small forest-steppe clearing in the forest, SRB obs. + photo (DJP det.); Bratislava – Podunajské Biskupice, Kopáč Island, PR Kopáčsky ostrov, ca. 48°5'41.97"N, 17°9'43.14"E, 132 m a.s.l., 13.vi.2006, 1 ♀, Malaise trap, MOB leg., coll. VKS; 30.v.2016, 2 ♂♂, and 3 ♀♀ FSLG after sunset, together with ca. 15 spec. of Od. armiger, EJB and RHB obs.; 7.vi.2016, 1 spec. FSLG after sunset, MSB obs.; 23.vi.2016, 2 ♂♂ FSLG at 21.32 and 21.38 CEST, MSB obs.; 1.vii.2016, 2 ♂♂ FSLG at 21.27 and 21.43 CEST, MSB obs.; 19.vii.2016, 4 spec. FSLG after sunset, EJB and JKB obs.; 20.vii.2016, 7 spec. FSLG after sunset, EJB and JKB obs.; ca. 48°5'39.5"N, 17°9'42.3"E, and 48°5'43.8"N, 17°9'30.4"E, 14.viii.2016, 7 spec. FSLG after sunset, DJP, ITV and VKB obs.; ca. 48°5'45.8"N, 17°9'41.9"E, 19.v.2018, 1 ♂ and 1 ♀ FSLG after sunset, MRV and VMP obs.; 9.vi.2018, 7 ♂♂ and 9 ♀♀ FSLG after sunset, together with ca. 20 spec. of Od. armiger, JHP, MRV and VMP obs.; 14.vii.2018, 1 ♂ and 4 ♀♀ FSLG after sunset, JRC and MRV obs.
Bratislava, Kopáč Island, PP Panský diel (Slovakia) in 2016 (before its conversion due to inappropriate conservation management) A–D site details (A photographed by Dalibor Všianský B–D photographed by Ilja Trojan) E push-up of B. unicornis (photograph by Milan Štrba) F male excavated from its burrow (photograph by Dalibor Všianský).
7869–7969: “Štefánikovce” [= Rovinka near Dunajská Lužná], ca. 130 m a.s.l., May 1949, tens of spec. observed during the day sitting on the tops of the grass blades above the water on a flooded steppe meadow (after the flood), Josef Marvan obs., 2 spec. (♂ and ♀) leg., coll. IMP.
7969: Bratislava – Čunovo, PR Ostrovné lúčky (Fig.
Data on flights of adults of B. unicornis at the locality of PR Ostrovné lúčky (for abbreviations see Table
Slovakia, Bratislava – Čunovo, PR Ostrovné lúčky | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
25.vii.2016 | 10 (6/4) | 21.10 | 21.37 | 20.37 | 33 min | 60 min | 27 min | 24 °C | before rain, no wind |
29.vii.2016 | 3 (2/1) | 21.11 | 21.25 | 20.31 | 42 min | 56 min | 14 min | 22 °C | no wind |
n | 13 (8/5) | average | 38 min | 58 min | 21 min | 23 °C |
Data on flights of adults of B. unicornis at the locality of Kalinkovská lesostep (for abbreviations see Table
Slovakia, Kalinkovo, Kalinkovská lesostep | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
27.vii.2016 | 3 (1/2) | 21.04 | 21.15 | 20.34 | 30 min | 41 min | 11 min | 22 °C | no wind |
7272: Višňové, Čachtický hradný vrch hill, 22.v.1988, 1 ♂ dead on a forest-steppe slope, IPO leg., coll. DJP.
7572: Hlohovec env., Nová hora near Koplotovce, 48°28'24.98"N, 17°49'21.40"E, 260 m a.s.l., 12.iv.1988, 1 ♀, accidentally dug up while turning the soil in the garden, KPH leg., coll. VKS; Hlohovec env., Mlynárska hora near Koplotovce, 48°28'6.51"N, 17°49'29.04"E, 235 m a.s.l., 4.vi.2021, 1 ♀ at light at 21.10 CEST (= 27 minutes after sunset), TSH obs.
7772: Šoporňa, [ca. 122 m a.s.l.], July 1952, 1 ♂, Kotek leg., coll. MHKC.
7373: Modrovka, [ca. 170 m a.s.l.], 15.vii.1979, 1 ♀, Mrklovský leg., coll. Ladislav Bojčuk deposited in MHKC.
7074: Trenčín – Zlatovce [env.], June 1926, [Rudolf] Čepelák leg., 1 ♂ and 1 ♀ in coll. Josef Gottwald deposited in
7174: “Trencsen Ungarn” [= Hungary, Trenčín], undated, 1 ♂ and 1 ♀ (ex coll. Engelbert Pawlik) in coll.
7274: [Považský Inovec Mts], Inovec [hill env.], 1 ♂, [Ladislav] Krejcárek [leg.], coll. Josef Gottwald deposited in
7674: Nitra [env.], 1950, no other data, 1 ♂ in coll. MHKC.
8177: Štúrovo env., Belianské kopce hills, “Hegyfarok” [= Modrý vrch], 47°49'8.09"N, 18°39'32.4"E, ca. 150 m a.s.l., 20.viii.2005, 1 spec., at light after midnight, 1.ix.2005, 1 spec., at light after midnight, 14.vi.2006, 2 spec., at light, 15.vi.2006, 1 spec., at light, 15.vi.2007, 1 spec., at light, 29.vii.2008, 2 spec., at light, 30.vii.2008, 1 spec., VVO obs.; Modrý vrch, PR Vŕšok, 47°49'6"N, 18°39'33"E, ca. 150 m a.s.l., 22.v.2014, 1 ♂, at light, OSO obs.; 47°49'13.5"N, 18°39'21.5"E, ca. 195 m a.s.l., 4.vi.2015, 1 ♀ at UV light at 21.30–0.30 CEST and 1 ♀ at UV light (the same trap) at 1.15 CEST (5.vi.2015), OSO obs. (moreover ca. 50 spec. of Och. integriceps Semenov, 1891 in the light traps were observed); 6.vi.2015, 1 ♂ FSLG after sunset, anonymous observer from the Czech Republic obs. (moreover 1 spec. of Och. integriceps in the light trap was observed); 47°49'9.54"N, 18°39'26.4"E, ca. 170 m a.s.l., 27.v.2015, 1 spec. FSLG after sunset, and 1 spec. at light, two anonymous observers from the Czech Republic obs.
8078: Zalaba, 47°58'8.8"N, 18°42'29.2"E, ca. 150 m a.s.l., June 1975, 1 spec. crawling on the ground on a sandy slope sparsely covered with black locust trees (Robinia pseudoacacia) at ca. 19.00 CEST, JAH.
8178: Bajtava, 16.vi.2006, 1 spec., at light, VVO obs.; Kamenica nad Hronom, Čierna hora hill (Fig.
Data on flights of adults of B. unicornis at the locality of Čierná hora hill (for abbreviations see Table
Slovakia, Kamenica nad Hronom env., Čierna hora hill | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
6.vi.2010 | 5 (4/1) | 21.30 | 21.45 | 20.38 | 52 min | 67 min | 15 min | - | after the floods |
7.vi.2010 | 16 (10/6) | 21.23 | 21.55 | 20.39 | 44 min | 76 min | 32 min | - | - |
4.viii.2011 | 17 | 20.50 | 21.20 | 20.18 | 32 min | 62 min | 30 min | 18 °C | after ca. 10 days of persistent rainfall, ca. 2 hours after the rain has ceased, vegetation heavily soaked, no wind |
5.viii.2011 | 10 | 20.50 | 21.10 | 20.16 | 34 min | 54 min | 20 min | 20 °C | almost no wind |
6.viii.2011 | 8 | 20.40 | 21.00 | 20.15 | 25 min | 45 min | 20 min | 23 °C | almost no wind |
9.viii.2011 | 15 | 20.45 | 21.05 | 20.10 | 35 min | 55 min | 20 min | 17 °C | almost no wind |
11.viii.2011 | 14 | 20.40 | 21.05 | 20.07 | 33 min | 58 min | 25 min | 18 °C | almost no wind |
12.viii.2011 | 9 | 20.40 | 21.05 | 20.05 | 35 min | 60 min | 25 min | 18 °C | almost no wind |
13.viii.2011 | 16 | 20.30 | 21.05 | 20.03 | 27 min | 62 min | 35 min | 19 °C | almost no wind |
16.viii.2011 | 2 (1/1) | 20.35 | 21.05 | 19.58 | 37 min | 67 min | 30 min | 22 °C | dry, almost no wind |
7.vi.2013 | 22 (13/9) | 21.20 | 21.45 | 20.39 | 41 min | 66 min | 25 min | 18 °C | almost no wind |
8.vi.2013 | 15 (10/5) | 21.17 | 21.42 | 20.40 | 37 min | 62 min | 25 min | 20 °C | light air to light breeze |
12.vi.2013 | 15 | 21.20 | 21.35 | 20.43 | 37 min | 52 min | 15 min | 18 °C | soil heavily saturated with water after rain, soaked vegetation |
15.vi.2013 | 10 | 21.20 | 21.45 | 20.44 | 36 min | 61 min | 25 min | - | - |
3.ix.2014 | 8 (5/3) | 19.52 | 20.07 | 19.24 | 28 min | 43 min | 15 min | 20 °C | light air to gentle breeze |
4.ix.2014 | 13 (8/5) | 19.51 | 20.16 | 19.22 | 29 min | 54 min | 25 min | 22 °C | almost no wind |
5.ix.2014 | 4 (2/2) | 19.47 | 20.04 | 19.20 | 27 min | 44 min | 17 min | 23 °C | almost no wind |
9.ix.2014 | 2 (2/-) | 19.43 | 19.47 | 19.12 | 31 min | 35 min | 4 min | 22 °C | almost no wind |
5.vi.2015 | 2 (2/-) | 21.22 | 21.27 | 20.38 | 44 min | 49 min | 5 min | 22 °C | almost no wind |
6.vi.2015 | 3 (2/1) | 21.24 | 21.47 | 20.39 | 46 min | 68 min | 27 min | 23 °C | light air |
28.v.2016 | 6 | 21.05 | 21.10 | 20.31 | 34 min | 39 min | 5 min | - | - |
1.vi.2016 | ≈ 8 | 21.15 | 21.45 | 20.35 | 40 min | 70 min | 30 min | 24 °C | no wind, very wet after rain |
2.vi.2016 | ≈ 8 | 21.15 | 21.45 | 20.36 | 39 min | 69 min | 30 min | 24 °C | no wind, very wet after rain |
3.vi.2016 | ≈ 8 | 21.15 | 21.45 | 20.37 | 38 min | 68 min | 30 min | 24 °C | no wind, very wet after rain |
11.vi.2016 | 5 | 21.20 | 21.40 | 20.43 | 37 min | 57 min | 20 min | - | - |
n | 241 | average | 36 min | 58 min | 22 min | 21 °C |
8178–8278: “Parkaň” [= Štúrovo], [ca. 110 m a.s.l.], 1934, no other data, 1 ♀ in coll. MHKC; 1940, no other data, 1 ♂ in coll. MHKC; Štúrovo, July 1967, 1 ♀, collector unknown, coll. ASH.
8179: Chľaba env., Močiar (the site near the confluence of the Danube and Ipeľ rivers), 47°49'14.53"N, 18°50'52.72"E, 110 m a.s.l., 12.vi.2014, 1 ♂ FSLG at 21.40 CEST, together with 2 ♀♀ of Od. armiger, OSO obs.
7785: Cerová vrchovina Mts, Hajnáčka – Buková env., ca. 48°13'51.39"N, 19°58'26.32"E, 1.vi.1978, 1 ♂ crawling on the ground in the afternoon in sunlight, IJN leg. [storage of the specimen unknown]; Cerová vrchovina Mts, Hajnáčka – Buková env., “circular pasture under vággon” [ca. 48°13'43.88"N, 19°58'15.53"E], 23.vi.1990, 1 ♀ flying at 21.28 CEST, JVP leg., ex original coll. JVP, currently in coll.
Data on flights of adults of B. unicornis at the locality of Hajnáčka – Buková (for abbreviations see Table
Slovakia, Hajnáčka – Buková | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
27.v.2008 | 5 (-/5) | 21.10 | 21.35 | 20.26 | 44 min | 69 min | 25 min | 22 °C | first or second warm day after a colder period of persistent rainfall; no wind |
29.v.2008 | 4 (1/3) | 21.10 | 21.40 | 20.29 | 41 min | 71 min | 30 min | 21 °C | newly hatched, light-coloured ♂ (not included in these statistics) crawling on a t-shirt spread on the ground at the edge of the forest under an oak tree (Quercus cerris) at 19.55 CEST; during flights, no wind to light air |
4.vii.2009 | 7 (3/4) | 21.15 | 21.45 | 20.43 | 32 min | 62 min | 30 min | 22 °C | - |
5.vii.2009 | 3 (1/2) | 21.15 | 21.30 | 20.43 | 32 min | 47 min | 15 min | 20 °C | - |
28.v.2010 | 4 (1/3) | 21.10 | 21.25 | 20.26 | 44 min | 59 min | 15 min | - | ♂ flying fast and high (ca 1.5–1.8 m above the ground) around a pile of logs near the edge of the forest |
n (♂/♀) | 23 (6/17) | average | 39 min | 62 min | 23 min | 21 °C |
7785–7885: Cerová vrchovina Mts, Gemerský Jablonec, 48°12'0.44"N, 19°59'24.31"E, 250–265 m a.s.l., steppe hillside with shrubbery of Prunus spinosa and Rosa canina on the hilltop, 4.vii.2009, 1 ♂ a 3 ♀♀ FSLG at 21.30–21.50 CEST, FPT and JPH obs. (see Table
Data on flights of adults of B. unicornis at the locality of Gemerský Jablonec (for abbreviations see Table
Slovakia, Gemerský Jablonec | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
4.vii.2009 | 4 (1/3) | 21.30 | 21.50 | 20.43 | 47 min | 67 min | 20 min | - | - |
5.vii.2009 | 4 (3/1) | 21.30 | 21.50 | 20.43 | 47 min | 67 min | 20 min | - | - |
28.v.2010 | 4 (1/3) | 21.00 | 21.15 | 20.26 | 34 min | 49 min | 15 min | - | - |
n (♂/♀) | 12 (5/7) | average | 32 min | 65 min | 32 min |
7786: Cerová vrchovina Mts, Hostice – Katarínka env., 48°13'52.55"N, 20°5'0.82"E, 216 m a.s.l., small steppe hillside with rich low vegetation and shrubbery of Prunus spinosa, 6.vi.2010, 2 ♀♀, hovering on the spot ca. 20 cm above the ground at 21.15 and 21.30 CEST (25 spec. of Od. armiger and 4 spec. of Och. chrysomeloides were also observed at the site), DJP obs. (see Table
Data on flights of adults of B. unicornis at the locality of Hostice – Katarínka (for abbreviations see Table
Slovakia, Hostice – Katarínka | |||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
6.vi.2010 | 2 (-/2) | 21.15 | 21.30 | 20.35 | 40 min | 55 min | 15 min | - | - |
7489: Slovak Karst, “Rakaťa” [= Rakyta Cottage] env., 48°35'29.7"N, 20°34'01.45"E, ca. 540 m a.s.l., 5.vii.1988, 1 ♀ excavated from its burrow, DKP (for partial data on this record see
7390: Slovak Karst, Hrhov, E of Okrúhle hill, 48°36'48.83"N, 20°47'22.98"E, ca. 395 m a.s.l., 4.vii.1988, 2 ♂♂ excavated from their burrows, DKP (for partial data on this record see
Comment
Slovakia is the country with the largest number of individuals found, as well as with the second largest number of known localities where the species has been recorded (52 sites). In addition to the already known localities,
Published data
“Gallia”, no other data (
“Alsace”, no other data (
Grand Est, Bas-Rhin, Strasbourg, ca. 140 m a.s.l., “in coll. Dr Puton – Jules Bourgeois pers. comm.”, no other data (
? Auvergne-Rhône-Alpes, Savoie, Albertville, 19th century, no other data, 1 ♂ and 1 ♀ in coll. Perroud deposited in
Material examined
? “S. Frankreich” [= south of France], 1 ♂ and 1 ♀, “Coll. C. Felsche, Kauf 20, 1918”, coll. MTDG [locality probably mistaken].
Grand Est, “Alsatia” [= Alsace], no other data, 1 ♂ in coll. Antoine Boucomont deposited in
? Occitanie, “Francia, Montpellier”, 1918, 1 ♀, Lavagne [leg.], coll. Paolo Luigioni deposited in
Comment
For France, B. unicornis was first recorded by
Published data
Baden-Württemberg, Markgräflerland, Neuenburg am Rhein – Grißheim, “Grißheimer Trockenaue”, [ca. 47°52'18.3"N, 7°33'55.5"E, ca. 210 m a.s.l.], 2.vi.1967, 1 ♂, at light, Hans Messmer leg., photo + coll. Richard Disch (
Bavaria (Bayern), “Bavaria”, no other data (
Material examined and new observations
“Germ.” [= Germany], no other data, 1 ♂ and 1 ♀ in coll. ZINR, 1 ♀ in coll. Karel Mazura deposited in
“Germania” [= Germany], no other data, 1 ♂ in coll. Georg Frey deposited in
“Germania mer.”, no other data, 1 spec in coll.
Baden-Württemberg, Bruchsal – Untergrombach, Michaelsberg and Habichtsbuckel Nature Reserve, ca. 49°5'32"N, 8°34'13"E, 200–220 m a.s.l., 3.vii.2021, 1 ♀, light trap, FTK and TBK obs., 4.vii.–5.viii.2021, 17 ♂♂ and 11 ♀♀ FSLG after sunset, together with more spec. of Od. armiger and Och. chrysomeloides, FTK and TBK obs. (5 spec. leg., coll. FTK, TBK and
Bavaria (Bayern), Upper Bavaria (Oberbayern), Ingolstadt, [ca. 370 m a.s.l.], 9.ix.[18]92, 1 spec. Dr K[arl] Daniel [leg.], “Fundortverwechslung” [= locality mistaken], coll.
Comment
In addition to old records from the late 18th and the first half of the 19th centuries from Bavaria, only one record from Baden from 1967 and two records from Bavaria in 1946 and 1954 were known from Germany. Daniel’s specimen from Ingolstadt and Kulzer’s specimen from Munich are questionable because they bear the labels added later of “Fundortverwechslung” (= locality mistaken). The new records presented from Baden represent the first known data on the species’ occurrence in Germany after 54 years (see also Fig.
Findings of B. unicornis A, B Germany, Bruchsal – Untergrombach, Michaelsberg and Habichtsbuckel Nature Reserve, 7.vii.2021 (photographs by Torsten Bittner) C, D Bulgaria, Dimovo env., 26.vi.2010 (photographs by Aleš Sedláček) E, F Bulgaria, Oreshak env., 6.vii.2020, (photographs by Maximilian Teodorescu).
Published data
Basel-Stadt (Kanton Basel-Stadt), Basel, undated, 1 spec., Ed. Bernoulli leg. (
? Republic and Canton of Ticino (Repubblica e Cantone Ticino), no other data, Villa [leg.] (
Material examined
Canton of Zürich (Kanton Zürich), “Tigurini” [= Zürich], undated, 2 ♀♀, collector unknown, “Mus. Drews.” [= Musaeum Drewseni, = ex coll. Christian Drewsen (1799–1896)], coll.
Comment
The two old records from Basel and Canton of Ticino were later considered questionable for the absence of any subsequent sightings (
Published data
“Italia”, no other data (
“Italia borealis”, no other data (
Piedmont (Piemonte), no other data (
Lombardy (Lombardia), no other data, (
Trentino-Alto Adige/Südtirol, “Tirol” [= probably Südtirol], [Stefano de] Bertolini leg. (
Veneto, no other data, (
Friuli Venezia Giulia, Provincia di Pordenone, Magredi del Cellina, Cordenons env., ca. 116 m a.s.l., 8.ix.2018, 1 ♂ and 1 ♀, dead on a path, 9.ix.2018, 1 ♂ and 1 ♀ in flight at 20.10–20.30 CEST (air temperature 21.5 °C, humidity 81%), 1 ♀ at actinic light at 21.15 CEST (air temperature 20 °C, humidity 96%), 10.ix.2018, 1 ♂ in flight at 20.15 CEST (air temperature 19 °C, humidity 78%), 12.x.2018, 1 ♀ in flight at 20.00 CEST, 15.v.2019, 1 ♂ and 1 ♀ in flight at 21.15 CEST (air temperature 14 °C, humidity 75%), 16.v.2019, 2 ♂♂ and 1 ♀ in flight at 21.00–21.15 CEST (air temperature 17 °C), 1 ♀ crawling on the ground at 21.20 CEST, 24.v.2019, 5 ♂♂ in flight at 21.20–21.35 CEST (air temperature 20 °C, humidity 70%), 26.v.2019, 1 ♂ in flight at 21.20–21.35 CEST (air temperature 20 °C, humidity 70%), 1.vi.2019, 1 ♂ and 2 ♀♀, in flight at 21.40 CEST (air temperature 20 °C, humidity 80%), 6.vi.2019, 2 ♂♂ and 1 ♀, in flight at 21.00–21.20 CEST (air temperature 22 °C, humidity 80%), 7.vi.2019, 2 ♂♂ in flight at 21.35 CEST (air temperature 22 °C, humidity 50%), Paolo Glerean and Gabriele Stefani obs. (
Tuscany (Toscana), no other data, coll. Dr L[ucas] von Heyden (
Material examined and new observations
“Ital.” [= Italy], no other data, 1 ♀ in coll. MNBG.
“Italia borealis”, 1 ♂, “ex coll. [Achille] Griffini”, no other data, coll.
“Italien” [= Italy], “coll. [Gustav] Kraatz”, no other data, 2 ♂♂ and 5 ♀♀ in coll.
“Italia”, undated, 1 ♂ and 1 ♀ (ex coll. Alexander Fry), coll.
“Italia, Sella [it is not clear whether it is a geographical name or the name of a person]”, no other data, 1 ♂ in coll.
Piedmont (Piemonte), “Pedem.”, [= Pedemontium, currently Piedmont], no other data, 1 ♂ in coll.
Lombardy (Lombardia), Provincia di Varese, Casorate Sempione, ca. 280 m a.s.l., October 1958, 1 ♂, at light, A. Bilardo leg., ex original coll. Giovanni Mariani, currently deposited in coll. RPM (for partial data on this record see
Comment
In the collection of Zdeněk Tesař deposited in SNMS, there is one specimen with the locality “Tirolis”, which may refer to the territory of South Tyrol (today Trentino-Alto Adige). Records from Sicily (
Published data
Mazovian Voivodeship (Województwo mazowieckie), Warsaw – Saska Kępa, 80–85 m a.s.l., undated, 2 spec., Antoni Waga leg. (
Opole Voivodeship (Województwo opolskie), Opole County, Złotniki, ca. 155 m a.s.l., undated, 1 ♀, Ludwik Fryderyk Hildt leg. (
Świętokrzyskie Voivodeship (Województwo świętokrzyskie), Kielce County, Chęciny, 1 spec., no other data, (
Lublin Voivodeship (Województwo lubelskie), Lublin env., no other data, Baumgarten leg. (
Material examined
Lesser Poland Voivodeship (Województwo małopolskie), Wadowice County (Powiat wadowicki), “Wadowice, Hal.” [= Hałyczyna or Galicja (Galicia), Wadowice, ca. 250–300 m a.s.l.], 1 ♀, undated, Smolik [leg.], DJP det., coll.
Świętokrzyskie Voivodeship (Województwo świętokrzyskie), Kielce County, “Góry Stokrzyskie [env.], Gałęzice, [Mt.] Góra Ostrówka” [currently the Ostrówka quarry, ca. 50°50'11.94"N, 20°24'46.38"E, ca. 250 m a.s.l.], July 1921, 1 ♂, J. Czarnocki [leg.], “Polonia, [coll.] Sz[ymon] Tenenbaum”, coll.
Comment
From Poland, only six records were known, which are summarised and specified by
Published data
Upper Austria (Oberösterreich), Linz, Scharlinz, ca. 250 m a.s.l., 25.v.1936, 1 ♂, [Johann] Wirthumer leg., coll. BZLA (
Lower Austria (Niederösterreich), no other data (
Vienna (Wien), no other data, coll. Dr Lucas von Heyden (
Burgenland, shore of Neusiedler See, plant materials amassed by flood water, several times (according to Sturm), with no further details (
Carinthia (Kärnten), Villach, Teufelsgraben, 1 spec. with no other data (
Styria (Steiermark), Grazer Bergland, Hörgas [near Gratwein-Straßengel], undated, 1 ♂ [10.6 mm], G[ustav] Wallaberger Sr. leg., coll.
Material examined
“Styria” [= Duchy of Styria, a territory that included the modern Austrian state of Styria and the Slovenian region of Lower Styria], 1858, 1 ♂, [Eduard Albert] Bielz [leg.], coll. BNMS.
Lower Austria (Niederösterreich), “Nied. Oesterr.” [= Niederösterreich], no other data, 1 ♂ in coll.
Vienna (Wien), “Wien” [= Vienna], no other data, 2 ♀♀ in coll. Vladimír Zoufal deposited in
Comment
In Austria, this species is known from six of the nine Austrian states. A recent attempt to rediscover the species at suitable sites along the Traun River in Upper Austria (
Published data
Western Transdanubia (Nyugat-Dunántúl), Vas County, “Molna-Szecsőd” [= Molnaszecsőd], 10.vi. [turn of the 19th and 20th century], ca. 180 m a.s.l., 1 spec. inside the digestive system of Cuculus canorus, Ernő Csiki obs. (
Central Transdanubia (Közép-Dunántúl), Komárom-Esztergom County, “Szőny” [a part of the current Komárom city], 6.viii.1901, 2 spec. inside the digestive system of Upupa epops, Ernő Csiki obs. (
Southern Transdanubia (Dél-Dunántúl), Somogy County, Fonyód, ca. 140 m a.s.l., undated, 1 ♂, Viktor Stiller leg., coll.
Central Hungary (Közép-Magyarország), Veszprém County, Pápa env., no other data (
Southern Great Plain (Dél-Alföld), Csongrád-Csanád County, Szeged – Kiskundorozsma env., Nagyszék, 16.–23.vi.1989, 1 spec., pitfall trap with ethylene glycol, Béla Gaskó leg., coll.
Northern Hungary (Észak-Magyarország), Heves County, Mátra Mts, Galyatető, 10.vii.1959, Sándor Szabó leg. (
Northern Great Plain (Észak-Alföld), Hajdú-Bihar County, “Debreczen” [= Debrecen] env., ca. 1860–1880, 1 spec., József Török leg. (
Material examined and new observations
Western Transdanubia (Nyugat-Dunántúl), Győr-Moson-Sopron County, Győr – Likócs env., ca. 47°42'52.5"N, 17°41'45"E, 2019, 115 m a.s.l., pitfall traps, no other data (see unpublished report for the Fertő-Hanság National Park Directorate,
Central Transdanubia (Közép-Dunántúl), Fejér County, Csór, ca. 150 m a.s.l., 21.v.2014, 1 ♀, at light on a steppe, MPK leg., coll. DCO; 28.v.2016, 1 ♂, at light, DVZ obs.; Fejér County, Csákberény, Bucka hill, 47°20'51.65"N, 18°21'35.32"E, 230 m a.s.l., 11.vi.1987, 1 spec., at light, CSB obs.; Fejér County, Gánt env., Köves-völgy [valley], 47°24'19.94"N, 18°22'47.67"E, 280 m a.s.l., 14.vi.2019, 1 ♀ flying after sunset, VSI obs.; Fejér County, Nagykarácsony, 46°52'49.4"N, 18°43'27.1"E, 150 m a.s.l., 2.vi.2021, 2 ♂♂ and 1 ♀ FSLG after sunset, 17–18 °C, light breeze, TDS obs.; Fejér County, Adony env., 47°5'17.2"N, 18°49'10.3"E, 120 m a.s.l., 1.vi.2021, 10 spec. FSLG after sunset, 11–15 °C, no wind, TDB and TDS obs.; Komárom-Esztergom County, Környe, no other data, 1 ♂ in coll.
Southern Transdanubia (Dél-Dunántúl), Somogy County, Balatonendréd, 46°50'52"N, 17°59'18"E, 174 m a.s.l., 11.v.1989, 1 ♀ excavated from its burrow together with 1 ♂ of Od. armiger, VRH; Somogy County, Ságvár, Jaba-völgy [valley], 46°49'28.29"N, 18°2'32.93"E, 180 m a.s.l., 25.ix.2017, 1 ♀ crawling on the ground, PFS obs. + photo (DJP det.); Somogy County, Balatonvilágos – Balatonaliga, 10.vi.1983, 1 spec., at light, SIB obs.; Baranya County, Zselic Mts, Mozsgó, ca. 150 m a.s.l., 27.vii.2017, 1 ♂, at light, MRM; Baranya County, Drávaszabolcs, 4/c Köztársaság tér Street, 45°48'20.95"N, 18°12'43.74"E, 91 m a.s.l., 28.vi.2020, 1 ♀ dead under the lamp, JST; Baranya County, Villányi-hegység Mts, Nagyharsány env., Szársomlyó hill, ca. 145 m a.s.l., 22.v.1977, 1 ♂, at light., AUP; Baranya County, Erdősmecske, ca. 240 m a.s.l., 18.viii.2012, 1 spec., 31.vii.2016, 1 spec., 27.v.2017, 1 spec., REE obs.
Central Hungary (Közép-Magyarország), Pest County, Zsámbék, June 2016, 1 ♀, students of Department of Zoology, Charles University, Prague leg., coll. DKP deposited in
Southern Great Plain (Dél-Alföld), Bács-Kiskun County, Kunpeszér env., Alsó-Peszéri-rétek, ca. 47°3'50.129"N, 19°17'57.59"E, 93 m a.s.l., 8.vi.1996, 2 ♂♂, 23.vi.1998, 1 spec., 10.vi.2002, 2 ♂♂ and 1 ♀, at light, AMK obs.; Bács-Kiskun County, Kunpeszér env., Peszéri-erdő forest, ca. 100 m a.s.l., 6.vi.1998, 6 spec., 21.vi.1999, 3 spec., 30.vi.1999, 2 ♂♂, 11.vii.1999, 1 spec., AMK obs.; 26.vi.2018, 2 spec., at light, REE obs.; 28.vi.2018, 3 spec., at light, REE and CVK obs.; 4.vii.2018, 4 spec., 5.vii.2018, 7 spec., at light, REE obs.; 6.vii.2018, 1 spec., CVK obs.; 9.vii.2020, 1 spec., 13.vii.2020, 1 spec., at light, REE obs.; 22.vii.2020, 1 spec., 29.vii.2020, 1 spec., 30.vii.2020, 5 spec., 31.vii.2020, 2 spec., 8.viii.2020, 1 spec., 16.viii.2020, 10 spec., 18.viii.2020, 7 spec., 19.viii.2020, 6 spec., 20.viii.2020, 8 spec., Botond Kozma obs.; Bács-Kiskun County, Kunadacs env., Hungarian meadow viper Conservation Centre, ca. 47°1'27.807"N, 19°17'21.286"E, ca. 100 m a.s.l., 29.vi.2018, 1 spec., Vadász Csaba obs.; Bács-Kiskun County, Kunadacs env., Hetvenholdas, ca. 47°0'56.34"N, 19°16'54.34"E, ca. 97 m a.s.l., 27.ix.2016, 1 ♀ FSLG after sunset, AMK obs.; Bács-Kiskun County, Kunadacs, Nagy-erdő forest, date not available [21st century], 1 spec. caught after sunset, AMK obs.; Bács-Kiskun County, Kunadacs, Peregi-dűlő, ca. 46°57'0.4"N, 19°17'29.4"E, 95 m a.s.l., 6.vi.2006, 1 spec., AMK obs.; Bács-Kiskun County, Kunadacs env., Szabadszállási-legelő, ca. 46°56'6"N, 19°18'15"E, 94 m a.s.l., 9.vi.2006, 1 spec., AMK obs.; Bács-Kiskun County, Páhi, Páhi-rétek, ca. 100 m a.s.l., 10.vii.2020, 3 spec., CBK and REE obs.; Bács-Kiskun County, Kiskunhalas env., pasture, 46°24'10.97"N, 19°30'33.08"E, 122 m a.s.l., 7.vi.2021, 2 ♂♂ and 1 ♀, at light just after sunset, together with 1 ♀ of Od. armiger, TKK obs. + photo (DJP det.); Bács-Kiskun County, Kecskemét – Hunyadiváros, 46°55'6.114"N, 19°42'43.794"E, 115 m a.s.l., 29.vi.2021, 1 ♀, at light, BCK and KVB obs. + photo (DJP det.).
Northern Hungary (Észak-Magyarország), Nógrád County, Kozárd, village area, 47°54'53.31"N, 19°37'7.07"E, 180 m a.s.l., 28.vii.2020, 1 ♀, at light (wall lamp of a residential house), KHE obs.; Nógrád County, Kozárd env., Majorsági-hegy hill, 47°54'59.87"N, 19°36'37.52"E, ca. 240 m a.s.l., 1.viii.2020, 1 ♀ flying after sunset, KHE obs.; Nógrád County, Kozárd env., Pohánka hill, 47°54'56.21"N, 19°37'29.88"E, 225 m a.s.l., 29.vii.2020, 1 ♂ and 1 ♀ flying after sunset, KHE obs.; Nógrád County, Bátonyterenye – Kisterenye, Váci Mihály Street, 48°0'32.22"N, 19°49'46.48"E, 190 m a.s.l., 28.v.1978, 1 ♂ FSLG after sunset, TKB and TKG obs.; Heves County, Tarnalelesz env., Pataji-far, 48°7'32.66"N, 20°9'32.12"E, 475 m a.s.l., 9.vi.2016, 1 ♂, in the grass during the day, shrubby edge of an oak forest (Quercus cerris), CBE obs.
Comment
Approximately one-third of the known localities of the species are located in Hungary. It is known here from 18 of the 19 counties. The first record from Hungary, without further details, is given by
Published data
Carniola (Kranjsko), “Carniolia, Bolboceras quadridens Fabr.”, undated, 1 ♂, Ferdinand Joseph Schmidt leg., coll. F. J. Schmidt deposited in
Styria (Štajerska), “Leonhard” [= Lenart v Slovenskih goricah], no other data, 1 spec. J. N. Spitzy leg. et coll. (
Comment
Although there are many localities suitable for the species in Slovenia, only five old records are known from this country. The species is most likely still present here, and the reason for the absence of new data is probably the low collecting activity of the local entomologists and/or the ignorance of appropriate monitoring methods for the species. Also, a recent attempt to rediscover the species in Slovenia (
Published data
Croatia proper (Središnja Hrvatska), Moslavina [a microregion between the rivers Lonja in the south and west, Česma in the north and Ilova in the east], no other data [19th century] (
Slavonia (Slavonija), Osijek-Baranja County, Osijek env., no other data [19th century], Vukas [leg.] (
Material examined
“Chorvatsko” [= Croatia in Czech language], 1 ♂, “ex. coll. E. Hachler”, no other data, coll.
Dalmatia (Dalmacija), “Dalmatia”, no other data, 1 ♀ in coll
Comment
In Croatia, the species is known only from four old records from the 19th century. The only recent record (Gradište) is given by
Published data
“Herzegovina”, no other data, 1 ♂ in coll.
Federation of Bosnia and Herzegovina (Federacija Bosne i Hercegovine), Zavidovići env., Gostović river valley, no other data, Károly Kendi leg. (
Federation of Bosnia and Herzegovina (Federacija Bosne i Hercegovine) or Republika Srpska (Република Српска), Babin potok [river], no other data, 1 spec. in coll. René Mikšić [currently deposited in
Comment
Only four old records from Bosnia and Herzegovina have been published. No recent findings are known.
Published data
Vojvodina (Војводина), Srem District (Сремски округ), Mt. Fruška gora (Фрушка гора), village of Vrdnik (Врдник), June 2016, 1 ♀, at light, collector unknown, coll. DKP deposited in
Belgrade District (Град Београд), Mala Ivanča (Мала Иванча) env., Grkovo (Грково), Trešnja Forest (Шума Трешња), 14.v.1986, 1 ♀ dug up beneath a hazel shrub together with Tuber fungi DPB leg. et coll. (
Southern and Eastern Serbia (Јужна и источна Србија), Bor District (Борски округ), Đerdap National Park (Национални парк Ђердап), 6 km WSW of Tekija (Текија), 27.–28.v.2014, 2 ♂♂ and 1 ♀, collector not specified, coll. DKP [deposited in
Material examined and new observations
Vojvodina (Војводина), Srem District (Сремски округ), Inđija (Инђија) env., Krčedin (Крчедин), 19.viii.2006, 2 ♂♂, at light, steppe meadow near the Danube river, LMN leg., coll. RSG and VVO; 45°10'04.5"N, 20°08'15.4"E, 98 m a.s.l., 1.vii.2013, 1 ♂, at light at 21.25 CEST, ZBB obs. + photo (for partial data on this record see
Findings of B. unicornis A Slovakia, Slanská Huta env., 24.vii.1972 (photograph by Zdeněk Laštůvka) B Serbia, Bukovac env., Beljevo hill, 27.v.2019 (photograph by Marko Šćiban) C Serbia, Leskovo env., 20.vi.2020, (photograph by Miloš Popović) D Ukraine, Semyhiria env., 3.vii.2020 (photograph by Dmytro Protopopov).
Southern and Eastern Serbia (Јужна и источна Србија), Bor District (Борски округ), Leskovo (Лесково) env., 44°18'17.28"N, 21°56'54.96"E, ca. 400 m a.s.l., 20.vi.2020, 1 ♂ crawling on the ground near the road at 18.42 CEST, MPN obs. + photo (Fig.
Data on flights of adults of B. unicornis at the locality of Tekija (for abbreviations see Table
Serbia, Tekija env. | |||||||||
date | n | BF | EF | S | S-BF | S-EF | DF | T | note |
27.v.2014 | 27 | 20.35 | 21.00 | 20.03 | 32 min | 57 min | 25 min | 24 °C | after a period of persistent rainfall |
28.v.2014 | 24 | 20.35 | 21.00 | 20.03 | 32 min | 57 min | 25 min | 24 °C | dtto |
12.vii.2014 | 9 | 20.45 | 21.15 | 20.16 | 29 min | 59 min | 30 min | 19 °C | ca 5 hours after the rain; no wind |
13.vii.2014 | 11 | 20.43 | 21.14 | 20.15 | 28 min | 59 min | 31 min | 22 °C | full moon, clear skies, light breeze, storm in the distance |
n | 71 | average | 30 min | 58 min | 28 min | 22 °C |
Data on flights of adults of B. unicornis at the locality of Cordenons env. (for abbreviations see Table
Italy, Cordenons env. ( |
|||||||||
---|---|---|---|---|---|---|---|---|---|
date | n (♂/♀) | BF | EF | S | S-BF | S-EF | DF | T | note |
9.ix.2018 | 2 (1/1) | 20.10 | 20.30 | 19.35 | 35 min | 55 min | 20 min | 21.5 °C | humidity 81% |
16.v.2019 | 3 (2/1) | 21.00 | 21.15 | 20.34 | 26 min | 41 min | 15 min | 17 °C | male crawling on the ground at 21.20 CEST |
24.v.2019 | 5 (5/-) | 21.20 | 21.35 | 20.43 | 37 min | 52 min | 15 min | 20 °C | humidity 70% |
6.vi.2019 | 3 (2/1) | 21.00 | 21.20 | 20.55 | 5 min | 25 min | 20 min | 22 °C | humidity 70% |
n (♂/♀) | 13 (10/3) | average | 26 min | 43 min | 18 min | 20 °C |
Comment
The known distribution of the species in Serbia was summarised by
Published data
Tirana County (Qarku i Tiranës), Sauk, 10.vi.1958, 1 spec., 10.–20.vi.1961, 1 spec., Xhelo Murraj leg. (
Comment
From Albania, only these records from two localities near Tirana have been published.
Published data
Crișana, Arad County, “Újarad” [= Arad – Aradul Nou], 28.iv.1907, 1 spec. inside the digestive system of Falco vespertinus, Ernő Csiki obs. (
Transylvania (Transilvania), Sălaj County, Zalău env., 3.viii.1973, 1 ♀, forest, collector unknown, coll. OHS (
Western Moldavia (Moldova Occidentală), Suceava County, “Mihoweny” [= Mihoveni], 1 ♂ with no other data (
Banat, Caraș-Severin County, “Gerník” [= Gârnic] env., 44°45'36.72"N, 21°46'29.48"E, 620 m a.s.l., 11.–13.vi.2016, ca. 14 spec. fyling low above the ground after sunset, together with tens of spec. of Od. armiger, air temperature 12–15 °C, JHH, JPP, JSU MSZ, MPV and PIL obs. (
Muntenia, Giurgiu County, Comana, no other data, Arnold Lucien Montandon leg., Jules Bourgeois det. (
Dobruja (Dobrogea), Tulcea County, Babadag [env.], [100–200 m a.s.l.], 1989–2000, no other data (
Material examined and new observations
Transylvania (Transilvania), Sibiu County, “Transsylv. Alpen” [= Transsilvanische Alpen (Carpații Meridionali)], “R.Turm Paſs” [= Roter-Turm-Pass (Pasul Turnu Roșu)], 350–450 m a.s.l., 1917, 1 ♂ and 1 ♀, Dr Maertens [leg.], coll. MNBG; Cluj County, Suatu, ca. 46°46'39"N, 23°58'24"E, ca. 365 m a.s.l., August 1997, 1 ♀, at light, steppe hillside with sparsely scattered oak trees, ARC (for incomplete data on this record see
Western Moldavia (Moldova Occidentală), Bacău County, Comănești (ca. 46°25'38.5"N, 26°26'31.1"E), July 2004, 1 ♂, dead inside the collector’s house (probably attracted by the light), APC; 31.vii.2010, 1 ♀, at light, APC; 8.viii.2011, 1 ♂, Barber pitfall trap, APC leg., coll. CMI; Iași County, Hârlău env., Pîrcovaci env., 47°28'28.29"N, 26°47'22.17"E, 240 m a.s.l., 24.vi.2021, 1 ♀, LHI obs. + photo (DJP det.); Iași County, Iași – Rediu, Iazul Tăutești, 47°13'33.4"N, 27°28'06.7"E, 120 m a.s.l., 28.vii.2021, 3 ♀♀, at light, MJR leg., coll. PKG; Iași County, Iași – Miroslava, Valea lui David, 47°11'38"N, 27°28'2.114"E, ca. 90 m a.s.l., 9.vii.2021, 2 ♀♀, together with 1 ♂ of Od. armiger, LHI obs. + photo (DJP det.); Iași County, Bârnova Forest (ca. 47°00'37.4"N, 27°33'32.8"E), 4.vii.2005, 1 ♂, found accidentally on the ground during the day, LFI leg., coll. CUIR (for incomplete data on this record see
Banat, Caraș-Severin County, Svatá Helena (Sfânta Elena) env., Kulhavá skála hill env., 44°42'11.47"N, 21°43'41.49"E, 357 m a.s.l., 1.vi.2012, 1 elytron on a path going through a pasture, BJN; Caraș-Severin County, Svatá Helena (Sfânta Elena), 44°40'29.8"N, 21°42'35"E, 325 m a.s.l., 18.vi.2017, 1 ♀ FSLG after sunset, ZCP, 44°40'57.73"N, 21°42'19"E, 350 m a.s.l., 23.vi.2017, 4 spec. FSLG after sunset, ZCP obs. (1 ♂ leg. et coll.); Caraș-Severin County, Mehadia, undated [19th century], 1 ♀, “ex. coll. [Otto] Staudinger”, coll. MTDG; Mehedinți County, Tisové Údolí (Eibenthal), ca. 44°32'36.7"N, 22°10'20.4"E, ca. 420 m a.s.l., 28.v.2008, 1 ♂ flying slowly up to 0.5 m above a path crossing a forest-steppe meadow at 21.45 EEST (= 40 min after sunset), JKV.
Muntenia, “Bukarest” [= Bucharest], undated, 1 ♂, V[ladimír] Zoufal leg., coll. Vladimír Zoufal deposited in
Dobruja (Dobrogea), Tulcea County, Agighiol, 12.vi.1993, 1 ♀, Ioana Matache leg., coll. GANM; Tulcea County, Babadag [env.], [100–200 m a.s.l.], 20.vi.1958, 1 ♂, 20.vi.1968, 1 ♂, Nicolae Săvulescu leg., coll. GANM; 11.vii.1985, 1 ♂, at light (mercury-vapor lamp), foot of a forest-steppe loess hill, JHM leg., coll. VKS (for partial data on this record see
Comment
For Romania, which can be considered one of the countries at the centre of the species’ distribution, surprisingly small amounts of data have been published. New records from 22 Romanian localities are presented here.
Published data
Călărași District (Raionul Călărași), Bularda near Dereneu, ca. 165 m a.s.l., 16.vi.1931, 3 ♂♂ and 4 ♀♀, Nicolai Zubowsky leg., coll. N. Zubowsky deposited in NMCM (
Ialoveni District (Raionul Ialoveni), Dănceni, ca. 170 m a.s.l., 31.v.1929, 1 ♂, Nicolai Zubowsky leg., coll. N. Zubowsky deposited in NMCM (
City of Chișinău (Municipiul Chișinău), Chișinău, [ca. 100 m a.s.l.], 20.v. and 10.vii.[between 1900–1915], no other data (
Material examined
City of Chișinău (Municipiul Chișinău), Chișinău, 20.iv.1912, 1 ♀, Nicolai Zubowsky leg., Valeriu Derjanschi det., coll. Rodion Stepanov (box No. 10) deposited in NMCM.
Anenii Noi District (Raionul Anenii Noi), Hîrbovăț env., Hîrbovăț Forest, ca. 285 m a.s.l., June 1970, 1 ♂, Rodion Stepanov leg., Valeriu Derjanschi det., coll. R. Stepanov (box No. 28) deposited in
Comment
The first known record from Moldova (Chișinău) is mentioned by
Published data
“Gubernia podolska” [= Podolian Governorate (Подольская губерния) of the Russian Empire, now Ukraine] (
“Volhynien” [= Volhynian Governorate (Волынская губерния), a historical region of the Russian Empire that included almost the entire area of today’s Volyn Oblast, as well as the Rivne and Zhytomyr Oblasts, northern parts of the Ternopil and Khmelnytskyi Oblasts, parts of the Podlaskie and Lublin Voivodeships of Poland and Brest Region of Belarus], undated, 2 spec., prof. Bresser leg. (
? Ivano-Frankivsk Oblast (Івано-Франківська область), Chornohora (Чорногора) [mountain range], 9.viii.1939, 1 ♂, collector unknown, coll.
Ternopil Oblast (Тернопільська область), Ternopil Raion (Тернопільський район), Zboriv (Зборів), 19.viii.1937, 1 ♀; collector unknown, coll.
Chernivtsi Oblast (Чернівецька область), Bukovina (Буковина), Chernivtsi Raion (Чернівецький район), Chernivtsi (Чернівці), 4 spec. with no other data (
Vinnytsia Oblast (Вінницька область), Vinnytsia Raion (Вінницький район), Vinnytsia (Вінниця) env., August 1928, 1 ♂, caught in flight in the evening, collector unknown (
Odessa Oblast (Одеська область), Odessa Raion (Одеський район), Odessa (Одеса), 1827–1831, no other data (
Kyiv Oblast (Київська область), Kiyv (Київ), old town, May 1839, 4 spec. under a dead dog, June 1870, 1 spec. on a grassy path, Johann Heinrich Hochhuth leg. (
Cherkasy Oblast (Черкаська область), Zvenyhorodka Raion (Звенигородський район), Talne (Тальне), 1 ♂ with no other data (
Chernihiv Oblast (Чернігівська область), Novhorod-Siverskyi Raion (Новгород-Сіверський район), Novhorod-Siverskyi (Новгород-Сіверський) env., 51°59'N, 33°16'E, 18.vii.2003, 1 spec., I. V. Porokhniach leg., coll. GUNU (
Sumy Oblast (Сумська область), Shostka Raion (Шосткинський район), Matskove (Мацкове) env., ca. 51°28'48"N, 33°53'24"E, ca. 150 m a.s.l., 28.vii.2018, 1 spec., MZK (
Poltava Oblast (Полтавська область), Lubny Raion (Лубенський район), Lubny (Лубни), [ca. 160 m a.s.l.], July (year and nummber of specimens not specified), Kruhlik [leg.], coll. Provincial Museum of Poltava (
Dnipropetrovsk Oblast (Дніпропетровська область), Synelnykove Raion (Синельниківський район), Raivka (Раївка), 1.viii.2000, 1 spec., A. M. Sumarokov leg. (
Material examined and new observations
Zakarpattia Oblast (Закарпатська область), Mukachevo Raion (Мукачівський район), “Schönb Ungarn” [= Hungary, Schenborn (Шенборн)], [ca. 190 m a.s.l.], “coll. Kirsch”, undated, 1 ♂, coll. MTDG.
Ivano-Frankivsk Oblast (Івано-Франківська область), Kosiv Raion (Косівський район), Pistyn (Пістинь), [ca. 400 m a.s.l.], undated, 1 ♂,“A. St?kl” [the third letter is illegible] leg., coll. SMLU.
Ternopil Oblast (Тернопільська область), Chortkiv Raion (Чортківський район), “Torskie, pow[iat] Zaleszcz[yki]” [= Zalishchyky (Заліщики) Powiat, Torske (Торське)], [ca. 250 m a.s.l.], 27.vi.[19]33, 1 ♂, collector unknown, coll.
Chernivtsi Oblast (Чернівецька область), Bukovina (Буковина), Chernivtsi Raion (Чернівецький район), “Czernowitz” [= Chernivtsi (Чернівці)], no other data, 1 ♂ in coll. Georg Frey deposited in
Vinnytsia Oblast (Вінницька область), “Киевская г[уберния], Сквирский у[езд]” [= Kiev Governorate of the Russian Empire (disestablished 1925), Skvirsky Uyezd (incorrectly, it was actually Lipovetsky Uyezd), currently Vinnytsia Raion (Вінницький район)], “Ильинцы” [= Illintsi (Іллінці)], 14.vi.[year not specified], [ca. 215 m a.s.l.], 1 ♀, collector not specified, coll. ZINR (probably one of the two specimens mentioned by
Odessa Oblast (Одеська область), Bilhorod-Dnistrovskyi Raion (Білгород-Дністровський район), Karolino-Buhaz (Кароліно-Бугаз), Studentska (Студентська) railway station, ca. 46°9'56.34"N, 30°33'24.58"E, 22 m a.s.l., 15.vi.2017, 1 ♂ crawling on the ground, OKO.
Kyiv Oblast (Київська область), Bila Tserkva Raion (Білоцерківський район), “Hałajki, Kijow[ska] g[ubernia]” [= Kiev Governorate, Halaiky (Галайки)], [ca. 190 m a.s.l.], [probably 19th Century], no other data, 1 ♂ in coll.
Cherkasy Oblast (Черкаська область), Cherkasy Raion (Черкаський район), Kaniv (Канів) env., Kaniv Nature Reserve (Канівський природний заповідник), 49°43'12"N, 31°31'19"E, ca. 200 m a.s.l., 20.vi.1984, 6 spec. excavated from their burrows, steppe slope in a hornbeam forest, KVM and VGG leg., coll. MKY and MPGU.
Kirovohrad Oblast (Кіровоградська область), Oleksandriia Raion (Олександрійський район), Semyhiria (Семигір’я) env., 49°0'29.52"N, 32°54'21.24"E, 135 m a.s.l., 2.vii.2020, 1 ♂, at light, DPS obs. + photo (Fig.
Dnipropetrovsk Oblast (Дніпропетровська область), Dnipro Raion (Дніпровський район), Dnipro (Дніпро) [Dnipropetrovsk until 19 May 2016], Tunelna Balka tract (Тунельна балка) [the name of an area with oak forest in the southern part of the city, see Fig.
Comment
In the checklist of Ukraine (
Published data
Vratsa Province (Област Враца), Oryahovo (Оряхово), no other data (
Ruse Province (Област Русе), Vetovo (Ветово), no other data (
Razgrad Province (Област Разград), Razgrad (Разград) env., 16.v.1905, in the barracks area, number of specimens not specified, Andrey Markovich leg, 13.vi.1907, in the vineyards, number of specimens not specified, Andrey Markovich leg. (
Pleven Province (Област Плевен), Pleven (Плевен), April [year and collector not specified] (
Sofia Province (Софийска област), Gorna Malina (Горна Малина) – “ДЗС” [= Държавно земеделско стопанство, area of the State Farm], ca. 650 m a.s.l., 7.vii.1969, 2 ♂♂ and 1 ♀ excavated from the soil from a depth of ca. 10 cm on a pasture (northern slope), collector not specified (
Shumen Province (Област Шумен), Shumen (Шумен), ca. 200 m a.s.l., 1914, 2 ♂♂, Hanuš leg., coll. MYP (
Burgas Province (Област Бургас), “Michurin (Мичурин)” [= Tsarevo (Царево)], 29.–30.vi.1982, 1 ♂, at light., BSP (
Silistra Province (Област Силистра), Dulovo (Дулово) env., Karakuz forest (гора Каракуз) [note: the locality label states “Gora Kanagöl, Dulowsko”], 14.vi.1952, 1 ♀, P[encho Stefanov] Drenski leg., coll.
Material examined ans new observations
Vidin Province (Област Видин), Dimovo (Димово) env., steppe meadow near the Archar (Арчар) river, 43°45'28.7"N, 22°44'51.1"E, 110 m a.s.l., 26.vi.2010, 3 ♂♂ and 2 ♀♀ FSLG after sunset just before a storm, together with several spec. of Och. chrysomeloides, no wind, 26 °C, ASH (Fig.
Varna Province (Област Варна), Oreshak (Орешак) env., 43°17'50.67"N, 27°53'47.29"E, 300 m a.s.l., 6.vii.2020, 1 ♀ flying up to 0.5 m above the grass at ca. 22.00 EEST, forest-steppe clearing in an oak forest, MTM obs. + photo (Fig.
Comment
So far, only nine localities have been published for Bulgaria. This study presents new records from two additional sites.
Published data
Marmara Region (Marmara Bölgesi), Edirne Province, ca. 15 km E of Edirne [according to Walter Heinz, pers. comm., these were periodically flooded meadows on the banks of the Ebros River S of Edirne], 27.iii.[19]88, 1 ♀, WHS leg., coll. DKP deposited in
Published data
Aegean Region (Ege Bölgesi), Denizli Province, Denizli env., [Çürüksu River valley], “Goundely” [= Goncalı] [railway station env.], ca. 200 m a.s.l., May [19]26, 1 ♂, [Hans] Kulzer leg., coll.
The only two specimens of B. unicornis so far known from Asia A Turkey, Denizli env., [Çürüksu River valley], “Goundely” [= Goncalı] [railway station env.], May [19]26, [Hans] Kulzer leg., coll.
Material examined
Aegean Region (Ege Bölgesi), Aydın Province, [Büyük Menderes River valley], “Bereketli (Denizli)” [= Bereketli near Nazilli], ca. 80 m a.s.l., 5.vii.1965, 1 ♂, [Helio] Pierotti and [Antonello] Perissinotto leg., DJP det. (2021), coll. Helio Pierotti deposited in
Comment
Only the three records mentioned above are known for Turkey. A record from Osmaniye Province (Kadirli) reported by
Great Britain
Published data
East of England, Cambridgeshire, marshes between Peterborough and Wisbech, beginning of summer 1807, 1 ♂ and 1 ♀, plant materials alluviated by flooded River Nene, together with 2 ♂♂ and 3 ♀♀ of Od. armiger, William Skrimshire leg. (
Comment
Skrimshire’s record was probably adopted by several subsequent authors (e.g.,
Country not specified [probably Russia]
Material examined
“Kaukasus” [= Caucasus], locality and date not specified, 1 ♂, F. Bucknall [leg.], coll.
Comment
This specimen represents the first record of the species from the Caucasus region. Unfortunately, country is not specified (it could be Russian part of the area, but confusion of the locality label cannot be discounted).
Bolbelasmus unicornis was included in checklists, catalogues, and monographs dealing with the scarabaeoid fauna of several countries as follows: France (
No records are known from mainland Greece. Records from the Greek island of Crete (
Records from Cyprus (
All the records from the Soviet Union and Russia (e.g.,
The species has also been listed for Belarus, Montenegro, and the Republic of North Macedonia (
Figure
The northernmost known historical locality of B. unicornis is Warsaw (Poland), while the northernmost locality with a recent record is Novhorod Siverskyi (northern Ukraine). The southernmost historical locality is Denizli (southwestern Turkey), while the southernmost recent localities are Babin Kal (Serbia) and Oreshak (Bulgaria). The westernmost historical locality is Mulhouse (Alsace, France), while the westernmost recent localities are Bruchsal (Baden, Germany) and Lerma (Piedmont, Italy). The easternmost locality with a recent record of the species is Kocherezhky (Ukraine), which is also the easternmost known point of occurrence of the species.
Adults of B. unicornis spend most of their time underground. Above-ground activity is limited to short flight periods after sunset. Exceptionally, adults have been observed crawling on the ground during daylight hours (see Faunistic records). Flight statistics from each site are shown in Tables
More than 50 individuals of B. unicornis were excavated from their burrows from depths of 5–60 cm during daylight hours. The length and the shape of the burrows varied, with the male burrows often changing direction from vertical to horizontal, whereas the female burrows usually descended vertically, changing direction only slightly and often leading to depths greater than those of males. In the dry periods and at the end of the season, the beetles burrow to the depths of more than 50 cm (e.g., Tomáš Vendl obs., see Faunistic records). When excavating adults, sometimes two to three individuals were found in a single burrow, even of the same sex (e.g., two males; see Faunistic records and
As for the feeding habits of B. unicornis, in none of the observations made by the author was the burrow found to lead to the sporocarp of hypogeous fungus or to the mycorrhizal roots of a shrub or tree. Nothing that could be considered as their food was ever found close to the buried individuals. In contrast, the Hungarian researchers repeatedly excavated the beetles near Budapest from the immediate vicinity of sporocarps of the large-spored pea truffle Glomus macrocarpum, which were approximately the size of a fingernail, together with more specimens of Od. armiger (
Immature stages have only been described in a few species of bolboceratines (
Since we have virtually no knowledge of the diet of adults or larvae of B. unicornis, we can only speculate on what its diet consists of. Given the findings of Australian and Japanese researchers on related species, the likely food of adults appears to be hypogeous fungi (spores, hyphae, and sporocarps), while the food of larvae could be fine soil humus and/or soil bacteria. The previous hypothesis that the larvae of B. unicornis feed on sporocarps of hypogeous fungi (e.g.,
Life cycles have been documented for only a few bolboceratine species (
Assuming that the development of B. unicornis is similar, it is very likely that only adults, both old and newly emerged, overwinter. This assumption is supported by numerous records where both old, dark-coloured individuals with heavily abraded teeth of fore tibiae and fresh, pale-coloured individuals with sharp protibial teeth have been recorded at the beginning of the season (pers. obs.). Some bolboceratines have overlapping generations. For example, in the genus Odonteus, eggs, larvae, pupae, and adults have been observed together in a single branching burrow (
In the Pannonian Basin, the centre of the distribution of B. unicornis, adults are active from May to September, exceptionally as early as April and as late as October, with a significant peak in June and the first half of July (Fig.
Bolbelasmus unicornis is a stenotopic species, characteristic of Pannonian steppes, forest-steppes and sparse deciduous forests, especially dominated by oaks. It is often found on sandy substrate (e.g., surroundings of Győr, Kiskunság National Park), sandy-loess substrate (e.g., localities around the village of Čejč), gravelly-sandy-loess substrate (e.g., Dunajské luhy Protected Landscape Area), loess substrate (e.g., wider surroundings of Štúrovo, Cerová vrchovina Mts, Gödöllő Hills, Dniester Canyon National Nature Park) or limestone substrate (e.g., Slovak Karst, Aggtelek National Park, some localities around Budapest). Characteristic habitats are steppe or forest-steppe pastures at the edges of oak forests (e.g., Cerová vrchovina Mts, Slovak Karst), of oak-beech or beech forests (e.g., Slanské kopce hills, Đerdap National Park, some localities in Romania), oak-hornbeam or hornbeam forests (e.g., Pirot District of Serbia, Kaniv Nature Reserve in Ukraine), and of shrub zones (with e.g., Crataegus oxyacantha, Prunus spinosa, and Rosa canina). In the Kiskunság National Park it occurs in the Pannonic sand dune thicket (Junipero-Populetum albae) (
It seems that the fundamental requirement of the species is natural vegetation cover and soil undisturbed by agriculture. The same was noted by
Central European sites with substantial populations of B. unicornis are characterised by the occurrence of plant species such as Quercus spp., Crataegus oxyacantha, Prunus spinosa, Rosa canina, Festuca spp., Thymus spp., Orobanche spp., Scabiosa ochroleuca, Euphorbia cyparissias, Achillea millefolium, and Artemisia spp.
On describing the potential habitat of the related B. gallicus in southern France,
The Coleoptera that co-occur in Central European localities with B. unicornis include Lethrus apterus (Laxmann, 1770), Odonteus armiger, Ochodaeus chrysomeloides, O. integriceps, Gymnopleurus spp., Carabus montivagus Palliardi, 1825, C. scabriusculus Olivier, 1795, Capnodis tenebrionis (Linnaeus, 1761), Perotis lugubris (Fabricius, 1777), Ptosima undecimmaculata (Herbst, 1784), Sphenoptera spp., and Agrilus albogularis Gory, 1841 (observations by many collectors including the author).
The elevation of the sites where B. unicornis has been recorded varies between 20 and 800 m a.s.l. The average altitude of all known localities for which it could be at least approximately determined (n = 351) is 220 m a.s.l. It is therefore a species of lowland and lower hills.
The most effective method for monitoring this species is to capture adults during their flights after sunset with a net using a flashlight, preferably a headlamp, in suitable microhabitat. This collecting method was employed as early as the 1920s by Rudolf Čepelák (see
Alternatively, during the day, one may find the beetles in their burrows, which are indicated by small piles of excavated soil at the entrances (so called “push-ups”). These push-ups are similar to those of some large ground-nesting bees but the push-ups of the bees are conical, composed of uniformly loose soil about a central entrance, whereas the soil pushed up by the earth-borer beetles tends to form an irregular pile of lumps (Figs
Light trapping appears to be ineffective to capture this species (cf. also
Bolbelasmus unicornis is also difficult to find due to the fact that observable activity of adults (flights and digging underground tunnels with push-ups) occurs only after heavy rains, when the soil is damp and loose enough for the beetles to burrow easily. During the dry periods, when the soil is hard, and also in winter, the beetles are buried deeper in the ground and show no above-ground activity, making it very difficult to find them.
Below is the translation of a passage on B. unicornis from the diary of the excellent Czech coleopterist Rudolf Čepelák (1886–1972; Fig.
“Bolbelasmus unicornis Schrank
From 1.vi. to 15.vii. Zlatovce (Malá hora hill; Vinohrady), Ľutov (Pálenice hill), and certainly from Trenčín southwards everywhere on the south-eastern slopes.
The area of Malá hora is sparsely covered with grass, which reaches 40–50 cm in places. If it is a quiet evening (no wind), preferably without moonlight, at 9 pm they start flying about 20–30 cm above the ground. In my right hand I have a net with a handle about 10 cm long, not white but dark, and in my left hand a torch. I bend down and suddenly hear ‘zzzzz...’. If I feel it’s very close, I shine the torch and immediately catch it with the net. If I miss, it falls into the grass, where I would look for it vainly. It flies like a bee collecting pollen. If the grass is cut, they fly quite fast, like Geotrupes. The flights continue until ca. 9.30 pm. Sometimes I hear it on the ground, stridulating like a longhorn beetle. And then we see: the female digging a hole and the male removing away the excavated soil. I dug a hole to see what they were looking for, but all I found were healthy Ornithogalum or Gagea bulbs.”
I would like to thank all the colleagues and staff at the institutions who provided me with their data and/or photographs (for a list of these see Materials and methods). For providing faunistic data, photographs, literature that was not easily available, and/or consultations, I thank in particular Aleš Bezděk (Institute of Entomology, Biology Centre of the Czech Academy of Sciences, České Budějovice), Vítězslav Kubáň (Šlapanice), David Král (Department of Zoology, Faculty of Science, Charles University, Prague), † Ottó Merkl (Department of Zoology, Hungarian Natural History Museum, Budapest), Valentin Szénási and Sándor Bérces (Duna-Ipoly National Park Directorate, Budapest), Sándor Ugró and Róbert Enyedi (Kiskunság National Park directorate, Kecskemét), Krisztián Harmos (Bükk National Park Directorate, Eger), Tibor Danyik (Körös-Maros National Park Directorate, Szarvas), László Nádai (Budapest), Balázs Károlyi (Budapest), Tibor Kovács (Mátra Museum of the Hungarian Natural History Museum, Gyöngyös), Tibor Kisbenedek (Department of Natural History, Janus Pannonius Museum, Pécs), Yurii Ye. Skrylnyk (Research Institute of Forestry & Forest Melioration named after G. M. Vysotsky, Kharkiv), Yurii V. Kanarskyi and Yurii M. Geriak (Institute of Ecology of the Carpathians of the National Academy of Sciences of Ukraine, Lviv), Andrii I. Bachynskyi (Dniester Canyon National Nature Park, Zalishchyky), Bohdan M. Vasko (Department of biological methods of pest control, I. I. Schmalhausen Institute of Zoology of National Academy of Sciences of Ukraine, Kyiv), Valeriu Derjanschi and Svetlana Bacal (Institute of Zoology of the Academy of Sciences of Moldova, Chișinău), Luciano Ragozzino (Lerma), Paolo Glerean (Museo Friulano di Storia naturale, Udine), Alberto Ballerio (Brescia), Enrico Barbero (Dipartimento di Scienze della Vita e Biologia dei Sistemi, Università degli Studi di Torino), Roberto Poggi (Museo Civico di Storia Naturale “Giacomo Doria”, Genoa), Michele Zilioli (Laboratorio di entomologia, Museo Civico di Storia Naturale, Milan), Roberto Casalini (Museo Civico di Zoologia, Rome), Giuseppe M. Carpaneto (Dipartimento di Scienze, Università Roma Tre, Rome), Marcello Romano (Capaci), Petra Kranebitter (Museo di Scienze Naturali dell’Alto Adige, Bolzano), Daniel Lorenz (Franziskaner Gymnasium Bozen, Bolzano), Mauro Gobbi (Museo delle Scienze, Trento), Stefano Ziani (Meldola), Riccardo Pittino (Milan), Marcello Zorzetto (Museo Civico di Storia Naturale “Silvia Zenari”, Pordenone), Marco Uliana (Museo di Storia Naturale di Venezia Giancarlo Ligabue, Venice), Alain Drumont and Julien Lalanne (Royal Belgian Institute of Natural Sciences, Brussels), Antoine Mantilleri and Christophe Rivier (Muséum national d’Histoire naturelle, Paris), Henry Callot and Marie Meister (Musée Zoologique de l’Université et de la Ville de Strasbourg), Denis Keith (Muséum des Sciences Naturelles et de Préhistoire, Chartres), Christoph Germann (Biowissenschaftlichen Abteilung, Naturhistorisches Museum Basel), Guido Sabatinelli (Département d’entomologie, Muséum d’histoire naturelle, Genève), Michael Greeff (Department of Environmental Systems Science, Swiss Federal Institute of Technology, Zürich), Hannes Baur (Naturhistorisches Museum Bern, Switzerland), Harald Schillhammer (2. Zoologische Abteilung, Naturhistorisches Museum Wien, Vienna), Martin Schwarz (Entomologische Arbeitsgemeinschaft, Biologiezentrum Linz), Ulrike Hausl-Hofstätter (Abteilung Biowissenschaften – Zoologie, Universalmuseum Joanneum, Graz), Norbert Ruckenbauer (Naturkundliche Landessammlung, Museum Niederösterreich, Sankt Pölten), Andreas Eckelt (Tiroler Landesmuseum Ferdinandeum, Innsbruck), Helge Heimburg (Sammlungs- und Wissenschaftszentrum, Abteilung Zoologie, Landesmuseums für Kärnten, Klagenfurt am Wörthersee), Petr Zábranský (Institut für Forstentomologie, Forstpathologie und Forstschutz, Universität für Bodenkultur Wien, Vienna), Alexander Dostal (Vienna), Katja Neven (Staatliche Naturwissenschaftliche Sammlungen Bayerns, Zoologische Staatssammlung, Munich), Bernd Jaeger (Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Museum für Naturkunde, Berlin), Marianna Simões and Mandy Schröter (Senckenberg Deutsches Entomologisches Institut, Müncheberg), Olaf Jäger (Senckenberg Naturhistorische Sammlungen, Museum für Tierkunde, Dresden), Alexander Riedel (Staatliches Museum für Naturkunde Karlsruhe), Arnaud Faille and Aron Bellersheim (Staatliches Museum für Naturkunde Stuttgart), Dirk Ahrens (Zoologishes Forschungsmuseum Alexander Koenig, Bonn), Roland Schultz (Senckenberg Museum für Naturkunde Görlitz), Florian Theves (Landesanstalt für Umwelt Baden-Württemberg, Karlsruhe), Torsten Bittner (Staatsbetrieb Sachsenforst, NSG-Verwaltung Königsbrücker Heide/Gohrischheide Zeithain, Königsbrück), Oliver Hillert (Schöneiche bei Berlin), Walter Heinz (Schwanfeld), Przemysław D. Szymroszczyk (Museum and Institute of Zoology of the Polish Academy of Sciences, Warsaw), Melanya Stan and Ionuț Ș. Iorgu (“Grigore Antipa” National Museum of Natural History, Bucharest), Iulia Muntean (Brukenthal National Museum, Sibiu), Cornelia Chimişliu (Department of Natural Sciences, Museum of Oltenia Craiova), Teodora Toshova (Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Sofia), Pavel Stoev and Borislav Guéorguiev (National Museum of Natural History, Bulgarian Academy of Sciences, Sofia), Andrey V. Frolov (Zoological Institute of Russian Academy of Sciences, Saint Petersburg), Oscar Vorst (Naturalis Biodiversity Centre, Leiden), Aslak Kappel Hansen and Caroline A. Høegh-Guldberg (Natural History Museum of Denmark, University of Copenhagen, Copenhagen), Christoffer Fägerström (Biological Museum, Lund University, Lund), Hans Mejlon (Museum of Evolution, Uppsala), Sergei Tarasov, Jaakko Mattila and Pekka Malinen (Finnish Museum of Natural History LUOMUS, Helsinki), Dmitry Telnov (Department of Life Sciences, Natural History Museum, London), Darren J. Mann (Oxford University Museum of Natural History, Oxford), Roman Cséfalvay (Rohovce – Kyselica), Vladimír Janský (Slovak National Museum – Natural History Museum, Bratislava), Vladimír Smetana (Tekovské múzeum v Leviciach, Levice), Tomáš Jászay (Šariš Museum in Bardejov), Attila Balázs (Department of Forest Protection and Hunting, Faculty of Forestry and Wood Technology, Mendel University of Agriculture and Forestry, Brno), Miroslav Mikát (Museum of Eastern Bohemia in Hradec Králové, Hradec Králové), Jiří Procházka (Moravian Museum, Brno), Jan Walter (Západočeské muzeum v Plzni, Plzeň), Jiří Hájek (National Museum, Prague), Jindřich Roháček (Silesian Museum, Opava), Tomi Trilar (Slovenian Museum of Natural History, Ljubljana), Toni Koren (Association Hyla, Zagreb), Srećko Ćurčić (Institute of Zoology, Faculty of Biology, University of Belgrade, Belgrade), Yves Bousquet (Gatineau, Québec), Ondřej Sabol (Ostrava), and Aleš Sedláček (Hranice).
Many thanks also go to Peter Kurina (Gajary), Vlastimil Mihal (Přerov), Michael Balke (Zoologische Staatssammlung, Munich), Oleksandr O. Sukhenko (Dnipro), Dalibor Všianský (Brno), Maurizio Gigli (Roma), and Milan Štrba (Bratislava) for taking photographs of specimens or habitats of B. unicornis.
For comments on earlier drafts of the manuscript and for linguistic corrections I am very thankful to Terry F. Houston (Department of Terrestrial Zoology, Western Australian Museum, Perth). I also thank Michelle Rossini (Finnish Museum of Natural History LUOMUS, Helsinki), Andrey V. Frolov, and an anonymous reviewer for critical comments on the manuscript, and Nathalie Yonow for the perfect cooperation in the editorial processing of this paper.
Finally, I am grateful to my teacher of entomology from my youth, Vojtěch Jiříček (Prostějov), who showed me the beetle covered in this study for the first time when I was 13 years old, and to my long-time colleague and friend Ilja Trojan (Velké Němčice), thanks to whom I returned to entomology and was able to complete this study, which I dedicate to the memory of Rudolf Čepelák, the discoverer of the most efficient collecting method for Bolbelasmus unicornis. I thank his grandson Svätopluk Čepelák (Dlhá nad Váhom) for providing photographs of his grandfather Rudolf and an excerpt from his entomological diary.