A peer-reviewed open-access journal

Zookeys 909: 79-158 (2020)

doi: 10.3897/zookeys.909.46838 RESEARCH ARTICLE #$ZooKey S

http:/ / ZOO keys -pen soft. net Launched to accelerate biodiversity research

Revision of the Merodon serrulatus group
(Diptera, Syrphidae)

Ante Vuji¢c', Laura Likov', Snezana Radenkovi¢', Natasa Kodis Tubi¢',
Mihajla Djan', Anja Sebic', Celeste Pérez-Bafidén’, Anatolij Barkalov’,
Riistem Hayat’, Santos Rojo’, Andrijana Andri¢*, Gunilla Stahls®

| University of Novi Sad, Department of Biology and Ecology, Trg Dositeja Obradovica 2, Novi Sad, Serbia
2 Department of Environmental Sciences and Natural Resources, Faculty of Sciences III, Campus of San Vicen-
te, University of Alicante, Spain 3 Institute of Systematics and Ecology of Animals, Russian Academy of Sciences,
Siberian Branch, Novosibirsk, Russia 4 Department of Plant Protection, Faculty of Agriculture, Akdeniz Uni-
versity, Antalya, Turkey § University of Novi Sad, BioSense Institute, Dr Zorana Dindica 1, Novi Sad, Serbia
6 Zoology Unit, Finnish Museum of Natural History Luomus, University of Helsinki, Finland

Corresponding author: Laura Likov (laura.likov@dbe.uns.ac.rs)

Academic editor: X. Mengual | Received 27 September 2019 | Accepted 11 December 2019 | Published 5 February 2020
http://zoobank. org/22B7FF 16-D0A2-40F9-B20E-F7C6E0AF 1842

Citation: Vuji¢ A, Likov L, Radenkovié S, Kocis Tubi¢é N, Djan M, Sebié A, Pérez-Bafién C, Barkalov A, Hayat R,
Rojo S, Andrié A, Stahls G (2020) Revision of the Merodon serrulatus group (Diptera, Syrphidae). ZooKeys 909:
79-158. https://doi.org/10.3897/zookeys.909.46838

Abstract

The phytophagous hoverfly genus Merodon Meigen, 1803 (Diptera, Syrphidae), which comprises more
than 160 species distributed in Palaearctic and Afrotropical regions, can be differentiated into multiple
groups of species that harbor high levels of hidden diversity. In this work, the serrulatus species group of
Merodon is revised, providing an illustrated key to species, a detailed discussion on the taxonomic char-
acters and a morphological diagnosis, including also the first data about the preimaginal morphology of
this species group. The study includes characteristics of the 13 species of the M. serrulatus group, along
with the available distributional data. Moreover, descriptions are provided for seven new species, namely
M. defectus Vuji¢, Likov & Radenkovié sp. nov., M. disjunctus Vuji¢, Likov & Radenkovié sp. nov.,
medium Vuji¢, Likov & Radenkovié sp. nov., M. nigrocapillatus Vuji¢, Likov & Radenkovié sp. nov., !/.
nigropunctum Vujic, Likov & Radenkovié sp. nov., M/Z. opacus Vuji¢, Likov & Radenkovi¢ sp. nov., and
M. trianguloculus Vuji¢, Likov & Radenkovié sp. nov. In addition, the taxa M. serrulatus (Wiedemann in
Meigen, 1822), M. bequaerti Hurkmans, 1993, M. hirsutus Sack, 1913, M. kawamurae Matsumura, 1916,
M. sacki (Paramonov, 1936) and M. sophron Hurkmans, 1993 are redefined and redescribed. Following a

detailed study of the type material sourced from different entomological collections, the status of all avail-

Copyright Authors. 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.

80 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

able taxa related to M. serrulatus is revised and a new synonymy is proposed: M. tener Sack, 1913 syn. nov.
(junior synonym of M. serrulatus). The identity of M. trizonus (Szilady, 1940) could not be assessed as the
type specimens are lost. Thus, the name M. trizonus is considered as nomen dubium. The monophyly and
composition of this species group are assessed through Maximum Parsimony and Maximum Likelihood
analyses of the mitochondrial COI and nuclear 28S rRNA gene sequences.

Keywords

28S rRNA, COI, immature stages, lectotype, morphology, new species, new synonyms, taxonomy

Table of contents

Tinte@eltneti aia See 2 me Oe ae eee cree ae ee er ee ee 81
Materials aicitet ROO Sp aren sre atamet tissu tthe de pda ath OT aged ste ten sd ath Ted SONzets el 82
Mate rial ek ATIN TCAs crtasn ss eagnecereenes re he vetroube tigen si¥p ve ba pe eWPe tevatee tepPap seiesdnecuey danloee te 82
Tiisrrtcetionmall Acre niyiias exe ctecc vcs sous toa ete ast tica cave da gh Mee eatcse sath tone ese code Bocane roosts es 83
(Pexceinorde Stel. On ACHES i tstcee 8 Selec a Tie ete AB Snes kt ness a out Balogh pownenns 84
‘Tie stuchy-Ob pretmaginal monn Glogye.t a ced. te Minicee tennis auth che EnSabon ut cl 84
Wiech aired S15 cue Soot sesina Sires cag/ Ble nantes Stas Ue staat otvines evar ee Bh abel eatin ovateens 85
EROS ULES eect eee re ee ree tient te ic eee ee ee ee 86
VL CVO LON SOIUU TALIS ORONA 5 seas sndeen hi detin nba Hekate cee Bean tedebad ached alee Srkaneehaaaginde hdecdastapaeee 86
Taxonomy and nomenclature of the species belonging to the Merodon serrulatus
SPECICSCOTOM DL is hy liel coherence ta shiie vlebas ebidre iach ale vbacklrpiiled ns eis iuubacsTh plaebais vbelcts all
Merad on DEAUAEr IT AUER DNA NID. .sctinaratele sepstn Aieein cop tenes axeumivaronnased eae nereies 91
Merodon defectus Vuji¢, Likov & Radenkovié sp. nov. .....cseeeeseeeeeeeseeeseeeeeees 98
Merodon disjunctus Vuji¢, Likov & Radenkovié sp. nov.....csceseeeeeeeseeseereees 103
WAV Od OR UAIVSIEFLES SACI POF 8a ooo stot ols att esos ah ocain etn ck etdodalsuanecesecbeuserctica ston. 106
Merodon kawamurae Matsumura, 1916............ccccsssseseccosssecccessvsccccevsecccacevess 110
Merodon medium Vujic, Likov & Radenkovié sp. nov. ......sceseeeeeseceneeeneeeeees 13
Merodon nigrocapillatus Vujic, Likov & Radenkovié sp. nov........eeeeeeseeseees 116
Merodon nigropunctum Vujic, Likov & Radenkovié sp. nov. .....eeeeeeeeseeeeees bhp
Merodon opacus Vuji¢é, Likov & Radenkovié sp. nov.......eeeseeseeseeeeeseeeseeseees 121
Merodonsackit Patain@niove LIS GO) iceeoccsedeankss oc hacsplandistoces tunes tepccededaoeeelodeed 128
Merodon serrulatus (Wiedemann in Meigen, 1822)... .eseeeeseeseeeeeseeeseeeeees 129
Merodonsophron Aurkimarisn 993) x. cvcisecuisnesenese lemeravecncontderteersededecnessecstet 135
Merodon trianguloculus Vuji¢, Likov & Radenkovié sp. nov. ......ceeeeeeeeeeseees 137
Merodon trizonus (Szilady, 1940) nomen AUbium ...eecccecccssecssscessseeststesetseeen 140
Key to the Merodon serrulatus species SOUP .......sssssesssesseseceseeseeesecneseaeeeeeseeaees 140
IMIGLECUIAE MN CLEDCG. seni eto sde tt anise eeteale uni eks eter ace der Marmuwarenst 143
TEVISGUSSTOR Es. leadh asthe tas ctioanl eee an subtnenu aust tnaesch Vahpitats betes fut atabedloacot edad. ence Raden 145
Taxon delimitation andsinteerative taAxOmonay sc. See top teestacacencoecthe sae aay seeet verona 145
TYR ATURE SEAS tessa sc tsnsuge cytaeaaasaescvensg abba te hpdiess teeta cahaaedethledetadbeegearwatan cased 146

istry bution ali cssPeCle NAIVET SILVA: is. hatin acebeie yeosladiasleenqeatsetensiedeecsneteestecoqeal 147

Revision of the Merodon serrulatus group 81

NCR OW EC CTTETIEG ansioun,.c.Waceanicns ole nissnl ai agearohsedD Juadlin wether opus obs edutuaabingedenas 148
FRG LOCI CES ech antes leer TI sale i et Rn Aaa OE lM, A Aeon Oe 149
Supplemientaty: matetial sls yresc nse A naemaneasscarennmsntneet braserau seh Rees aie Releases h55
SU p le MiGhca ys Md Abe MAINS 5.5, Meeeies 05,6 OB ofa he meet A MCh Se eek nat etre kag alate 155
Slop lemGiiaty inate mercer MNP ee a LIE a Rh en, dere 156
Supplentenwbaty arate rial At ee ees 5 ee es, eee ee ee 156
Slip wle trie meat yer Aerial wo) erie te: sat Bu. Mee eee a See ee, 8 Bese 157
Supp lementatysriate ral@ Wns cu Oi a doit e ler Baer ah 157
SU LENT Mba oT AO LIAL rca. las teats Pa tes eptecac hoe O as or tee Maccecse cate nea eee eataceer 158
Supplemeritary material’ 2.5 ticesseus souies, consent cesgens soetsge pees et eee sees Tae catees 158
Introduction

The phytophagous hoverfly genus Merodon Meigen, 1803 contains more than 160 spe-
cies distributed across the Palaearctic and Afrotropical regions (Stahls et al. 2009). Adults
mimic bees and bumblebees (Hymenoptera: Apidae) and feed on pollen and nectar from
early spring to autumn (Hurkmans 1993; Speight 2018). Based on the immature stages
of Merodon found to date, underground storage organs (bulbs, corms and rhizomes) of
geophytes of the families Asparagaceae, Amaryllidaceae and Iridaceae are larval micro-
habitats of this taxon and phytophagy is its feeding mode (Ricarte et al. 2017; Preradovic
et al. 2018). The immature stages of only eight Merodon species have been described
to date (Heiss 1938; Stuckenberg 1956; Ricarte et al. 2008, 2017; Andrié et al. 2014;
Preradovié et al. 2018) and a detailed literature review on the immature stages of Mero-
don, including host plants, has recently been published by Ricarte et al. (2017).

The taxonomic status and identification of many Merodon species requires further
investigation, as the genus contains a high number of species groups consisting of
morphologically cryptic taxa with very subtle morphological differences. In various
recent publications, an integrative taxonomic approach combining morphological and
molecular information has been adopted and resulted useful in resolving taxonomic
ambiguities in hoverflies, e.g., in Merodon equestris species complex (Marcos-Garcia et
al. 2011), Merodon avidus complex (Popovié et al. 2015; Acanski et al. 2016), Merodon
aureus species group (Sasié et al. 2016), genus Chrysotoxum Meigen, 1803 (Nedeljkovi¢
et al. 2013, 2015) and Melanostoma Schiner, 1860 (Haarto and Stahls 2014).

Most recent publications pertaining to the genus Merodon have focused on par-
ticular species groups, within which the authors delimited individual species (Vuji¢
et al. 2012, 2013, 2015; Acanski et al. 2016; Sagié et al. 2016; Veselié et al. 2017:
Kocis Tubi¢ et al. 2018; Radenkovié et al. 2018a). The Merodon avidus-nigritarsis
lineage was confirmed as one of four main lineages in the genus Merodon, along-
side with three other lineages: albifrons+desuturinus, aureus and natans. Likov et
al. (2019) presented a phylogenetic inference where the Merodon avidus-nigritarsis
lineage was resolved in a similar way as in the studies by Sasié et al. (2016) and
Radenkovié et al. (2018b). In the same study, Likov et al. (2019) divided the

82 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

avidus-nigritarsis lineage into 10 species groups (namely M. aberrans, M. aurifer, M.
avidus, M. clavipes, M. fulcratus, M. italicus, M. nigritarsis, M. pruni, M. serrulatus,
and M. tarsatus groups), and five species were not included in any of these species
groups (i.e., M. clunipes Sack, 1913, M. crassifemoris Paramonov, 1925, M. eumerusi
Vuji¢, Radenkovié & Likov, 2019, M. murinus Sack, 1913, and M. ottomanus Hurk-
mans, 1993).

The Merodon serrulatus species group includes taxa with a characteristic basolateral
protrusion on the posterior surstyle lobe (Fig. 1). Based on recently published data,
this group contains six already described species, i.e., Merodon bequaerti Hurkmans,
1993, M. hirsutus Sack, 1913, M. kawamurae Matsumura, 1916, M. sacki (Paramonov,
1936), and M. serrulatus (Wiedemann in Meigen, 1822) (Likov et al. 2019).

In this study, we present a taxonomic review of the serrulatus species group based
on a detailed examination of material gathered as a part of our long-term field research
in the Palaearctic region, especially in the Mediterranean and the Middle East. Our
aims are 1. to review materials stored in several major entomological institutions and
private collections holding specimens of this group; 2. to define and describe the taxa
within the serrulatus species group, including new species; 3. to infer the phylogenetic
relationships among the members of this species group using mtDNA COI gene and
the 28S rRNA gene; and 4. to present the first data about the preimaginal morphology
of the M. serrulatus species group.

Materials and methods

Material examined

Most of the recently collected specimens were sampled by sweep net. Further speci-
mens of the Merodon serrulatus species group were sourced from collections deposited
in museums and universities which are listed below. Consisted total of 1,083 speci-
mens collected from 1837 to 2018 across 22 countries (i.e., Algeria, China, Croatia,
France, Greece, Israel, Italy, Kazakhstan, Kyrgyzstan, Libya, Montenegro, Morocco,
North Macedonia, Portugal, Russia, Spain, Syria, Tajikistan, Tunisia, Turkey, Turk-
menistan, and Uzbekistan) were studied for the present study.

The information on labels of the material examined is provided for each studied
specimen in the following order: country name, a bullet point (indicating the begin-
ning of a material citation), number and sex of specimen(s), locality data, geographi-
cal coordinates, altitude, collection date, collector(s) followed by “leg.”, institutional
acronym and specimen codes/unique identifiers (“to” indicates range). The specimens
are listed alphabetically by country and subsequently by increasing latitude (south to
north) within each country. In the quotations of the type specimens’ original label
data, double quotation marks were used to indicate separate labels, and the slash was
adopted to indicate a new line within a label, with additional details and interpreta-
tions provided in square brackets, where applicable.

Revision of the Merodon serrulatus group 83

Institutional acronyms

A. S. coll.
CEUA

D. D. coll.

EMIT
FSUNS

G. V. W. coll.

GLAHM

J. T. S. coll.

J. v. S. coll.

M. B. coll.

M. H. coll.

MAegean

MNHN
MZH

NBCN

NHMUK
NHMW

NMNH
NMPC
NMS

S. K. coll.

S. S. coll.
SIZK

SZMN

TAUI

Axel Ssymank collection, Achtberg, Germany (ssymanka@t-online.de)
Colecci6n Entomoldogica de la Universidad de Alicante, Alicante,
Spain

Dieter Doczkal collection, Munich, Germany (dieter.doczkal@gmail.
com)

Entomological Museum of Isparta, Isparta, Turkey

Faculty of Sciences, Department of Biology and Ecology, University of
Novi Sad, Novi Sad, Serbia

Guy Van de Weyer collection, Reet (Rumst), Belgium (guido.vandew-
eyer@skynet.be)

Hunterian Zoology Museum, University of Glasgow, Glasgow, UK
John T. Smit collection, Utrecht, the Netherlands (John.Smit@natura-
lis.nl)

Jeroen van Steenis collection, Amersfoort, the Netherlands (jvanstee-
nis! @gmail.com)

Miroslav Bartak collection, Prague, Czech Republic (bartak@af.czu.cz)
Martin Hauser collection, Sacramento, USA (martin.hauser@cdfa.
ca.gov)

The Melissotheque of the Aegean, University of the Aegean, Mytilene,
Greece

Muséum National d’ Histoire Naturelle, Paris, France

Finnish Museum of Natural History, University of Helsinki, Helsinki,
Finland

Naturalis Biodiversity Center [formerly known as the National Mu-
seum of Natural History (RMNH)], Leiden, The Netherlands
Natural History Museum, London, UK

Museum of Natural History (Naturhistorisches Museum Wien), Vi-
enna, Austria

The Department of Entomology, of the National Museum of Natural
History, Smithsonian Institution, Washington, DC, USA

National History Museum, Prague, Czech Republic

National Museums Scotland, Edinburgh, UK

Sakari Kerppola collection, Helsinki, Finland (sakari.kerppola@hel-
sinkinet.fi)

Stileyman Saribiyik collection, Kastamonu, Turkey

I.I. Schmalhausen Institute of Zoology of National Academy of Sci-
ences of Ukraine, Kiev, Ukraine

Siberian Zoological Museum of the Institute of Systematics and Ecol-
ogy of Animals, Siberian Branch of the Russian Academy of Sciences,
Novosibirsk, Russia

Tel Aviv University, Tel Aviv, Israel

84 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

WML World Museum Liverpool, Liverpool, UK

ZHMB Zoological (Zoologisches) Museum of the Humboldt University, Ber-
lin, Germany

ZMKU Zoological Museum, State University of Kiev, Kiev, Ukraine

ZMUC Zoological Museum, Natural History Museum of Denmark, Univer-

sity of Copenhagen, Copenhagen, Denmark

Taxonomic study of adults

The type material of all described species of the Merodon serrulatus species group were
studied, with the exception of the type material of Merodon trizonus (Szilady, 1940)
because the type specimens are lost.

To study the male genitalia, dry specimens were relaxed in a closed pot containing
water to ensure high humidity levels, and the genitalia were extracted using an insect
pin with a hooked tip. Genitalia were cleared by boiling them individually in tubes of
water-diluted KOH pellets for 3-5 minutes. This was followed by brief immersion in
acetic acid to neutralize the KOH, immersion in ethanol to remove the acid, and storage
in microvials containing glycerol. Specimens’ measurements were taken in dorsal view
with a micrometer and are presented as ranges. Body length was measured from the lu-
nule to the end of the abdomen. Drawings were made using a FSA 25 PE drawing tube
attached to a binocular microscope Leica MZ16. Specimens photographs were captured
by a Nikon D7100 camera connected to a personal computer, as well as a Leica DFC
320 digital camera attached to a Leica MZ16 binocular microscope. After photograph-
ing, CombineZ software (Hadley 2006) was used in order to create composite image
with an extended depth of field, created from the in-focus areas of each image.

Terminology adopted in the morphological descriptions follows Thompson (1999)
and, for male genitalia, Marcos-Garcia et al. (2007), while the term “fossette” follows
Doczkal and Pape (2009).

Localities were geo-referenced in Google Earth (Google Inc, California, USA,
https://www.google.com/earth; accessed on 10.02.2019). Geographic coordinates of
localities were represented in GenGIS (v 2.5.3) (Parks et al. 2013) in order to create
distributional maps.

The study of preimaginal morphology
Sampling

A targeted search for immature Merodon hoverflies was conducted in the chestnut
forest of Agiassos, Lesvos Island (Greece). Abundant population of M. serrulatus and
other Merodon species were found at this locality. Searches for larvae were carried out
during a field trip from February 27 to March 10 2006, as owing to their biological
cycle, these Merodon species would be in immature stages (larvae or pupae) during

Revision of the Merodon serrulatus group 85

this period. An area of ca. 3m’, where the presence of many bulb species and adults
of Merodon were reported the previous year was selected. The whole area was exca-
vated to a depth of approximately 20 cm and the soil was sieved searching for the
larvae. Only one larva (third larval stage) was found in the soil surrounding bulbs of
different plant genera, such as Fritillaria Tourn. ex L., Gagea Salisb., Muscari Miller,
and Ornithogalum L. This solitary larva was kept in a plastic container with the soil
in which it had been found at room temperature until it pupated two days later. The
adult of M. opacus sp. nov. emerged on 21 March 2006 after spending 17 days in
the pupal stage.

Morphological study

The cephalopharyngeal skeleton was removed from the antero-ventral margin of the
puparium using entomological pins. After dissection, the cephalopharyngeal skeleton
was soaked in 10% KOH and heated for 15 min in order to remove the remaining
tissues attached, after which it was soaked for a few minutes in acetic acid followed by
70% ethanol. Once the tissues had been cleared, the skeleton was preserved in glycerin.
Debris adhering to the puparial integument was removed with pins and brushes and
by placing the specimens in an ultrasonic cleaner for a few minutes. The cleaned speci-
men was mounted on stubs and was examined with a scanning electron microscope
(S3000N Hitachi) at 20 kV using variable-pressure (or low vacuum) mode, as this
technique allows a direct evaluation of the specimens without coating the samples with
gold. The stereomicroscope Olympus SZX16 (equipped with Olympus U-TVO.5XC-3
camera) was used for the examination and to capture images of the puparium (general
view) and the cephalopharyngeal skeleton. Dimension measurements (in mm) were
performed on preserved specimens using a Leica MZ9.5 binocular microscope.

The terminology for immature stages adopted here follows Rotheray (1993) and
Rotheray and Gilbert (1999), whereas certain characters of the cephalopharyngeal
skeleton are determined in line with Hartley (1963), and our morphological character
descriptions are based on Merodon puparia descriptions provided by other authors
(Heiss 1938; Stuckenberg 1956; Ricarte et al. 2008, 2017; Preradovié et al. 2018).
The studied material has been deposited at the University of Alicante, Spain (CEUA).

Molecular analysis

The specimens subjected to molecular analysis are presented in Supplementary file
8: Table 1. DNA voucher specimens were deposited in FSUNS, EMIT, SZMN, and
MZH. The genomic DNA of each specimen was extracted from two or three legs
using a slightly modified SDS extraction protocol (Chen et al. 2010). For this pur-
pose, the D2-3 region of the nuclear 28S ribosomal RNA gene and the mitochondrial
protein-coding cytochrome c oxidase subunit I (COI) gene were amplified. Primer
pair F2 and 3DR was used for the amplification of 28S rRNA gene region (Belshaw
et al. 2001), whereas C1-J-2183 (alias Jerry) and TL2-N-3014 (alias Pat) primer pair

86 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

(Simon et al. 1994) was chosen for 3'-end of COI gene, and for 5'-end COI gene, we
used LCO1490 and HCO2198 primer pair (Folmer et al. 1994). The PCR reactions
were carried out according to Kodis Tubié et al. (2018). The amplification products
were enzymatically purified by Exonuclease I and FastAP Thermosensitive Alkaline
Phosphatase enzymes (ThermoScientific, Lithuania) and sequenced using only forward
primers on an ABI3730x1 Genetic Analyzer (Applied Biosystems, Foster City, CA,
USA) at the Finnish Institute for Molecular Medicine (FIMM), Helsinki, Finland.

Data analysis

In order to establish the systematic position and composition of the Merodon serrulatus
group, samples representing the four main Merodon lineages were analyzed following the
approaches described by Sasié et al. (2016) and Radenkovié et al. (2018b), while two
further Merodontini species served as outgroups, i.e., Platynochaetus macquarti Loew,
1862 and Eumerus grandis Meigen, 1822 (see Supplementary file 8: Table 1 for GB
accession numbers of all analyzed species and outgroups). Alignment of the obtained
COI sequences was achieved using the Clustal W algorithm (Thompson et al. 1994)
implemented in BioEdit (Hall 1999), while rRNA 28S gene was aligned by the multiple
alignment using Fast Fourier Transform (MAFFT) program (Katoh et al. 2005, 2009),
version 7, which implements iterative refinement methods (Katoh and Standley 2013).
The E-INS-i strategy was chosen (Katoh et al. 2009). All sequences in the analyzed two-
gene dataset (concatenated COI and 28S rRNA gene sequences) were trimmed to equal
lengths. Phylogenetic tree construction was performed by conducting Maximum Parsi-
mony (MP) and Maximum Likelihood (ML) analyses. The parsimony analysis was con-
ducted using NONA (Goloboff 1999), spawned with the aid of ASADO, version 1.85
(Nixon 2008), using the heuristic search algorithm (settings: mult*1,000, hold/100,
max trees 100,000, TBR branch swapping). GTRGAMMA model was determined as
the best choice model for the analysed dataset using MEGA 7 (Kumar et al. 2016). The
dataset was divided into two partitions: COI gene and 28S rRNA gene. The ML tree
was constructed by RAxML 8.2.8 (Stamatakis 2014) using the CIPRES Science Gate-
way (Miller et al. 2010) and applying the general time-reversible (GTR) evolutionary
model with a gamma distribution (GTRGAMMA) (Rodriguez et al. 1990). Nodal sup-
port was estimated using nonparametric bootstrapping with 1,000 replicates for both
MP and ML trees, which were rooted on Platynochaetus macquarti.

Results
Merodon serrulatus group
Diagnosis. Member species of the Merodon serrulatus species group exhibit a distinc-

tive and characteristic basolateral protrusion (lateral hump) on the posterior surstyle

lobe (Fig. 1A, B: bp, 6: bp, 14C: bp). They are relatively large (11-15 mm) species with

Revision of the Merodon serrulatus group 87

Figure |. Merodon serrulatus male genitalia. A Epandrium, lateral view B Epandrium, ventral view C,
D Posterior surstyle lobe, lateral view E Hypandrium, lateral view F Part of hypandrium, ventral view G
Aedeagus, lateral view. Abbreviations: al—anterior surstyle lobe, bp—basolateral protrusion, c—cercus, |-lin-

gula, pl—posterior surstyle lobe, s—lateral sclerite of aedeagus. Scale bar: 0.2 mm.

a dark scutum and white microtrichose fasciae (at least in females) on the dark olive
brown terga 2—4 (as in Fig. 2); tergum 2 usually with a pair of reddish orange lateral
maculae. Antennae dark brown (as in Fig. 5).

88 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 2. Merodon serrulatus body parts, dorsal view. A thorax, male B thorax, female C abdomen, male

D abdomen, female. Scale bar: 2 mm.

Basoflagellomere 1.5—2.2 times as long as wide, usually obviously concave dor-
sally (as in Fig. 3A, D, G, J). Scutum covered with erect, usually yellow pile. Pile on
metasternum erect, and as long as those on metacoxa. Posterior part of mesocoxa bare,
without long pile. Legs mostly black, without spinae or other protuberances (as in Fig.
4). Metafemora incrassate (as on Fig. 4). Tarsi black dorsally and dark brown ventrally.
Abdomen elongated, narrow and tapering (as on Fig. 2), slightly longer than scutum

Revision of the Merodon serrulatus group 89

\
‘
\

Figure 3. Merodon serrulatus antenna. A outer side, lateral view, male (Spain) B inner side, lateral view,

ye

—

male (Spain) C dorsal view, male (Spain) D outer side, lateral view, female (Spain) E inner side, lateral
view, female (Spain) F dorsal view, female (Spain) G outer side, lateral view, male (Greece) H inner side,
lateral view, male (Greece) I dorsal view, male (Greece) J outer side, lateral view, male (Russia). Abbrevia-

tion: f-fossette. Scale bar: 1 mm.

and scutellum together. Male genitalia: apical part of anterior surstyle lobe more or less
of rhomboid or triangular in shape (as on Fig. 1A: al, C, D), covered with dense short
pile; posterior surstyle lobe oval with basolateral protrusion (lateral hump) (Fig. 1A,

90 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 4. Merodon serrulatus, lateral view. A, C metatrochanter, metafemur and metatibia B, D-F metaleg.
A-B male (Spain) C female (Spain) D male (France) E male (Greece) F male (Russia). Scale bar: 2 mm.

B: bp, 14C: bp); cercus rectangular, without prominences (Fig. 1A: c). Hypandrium
elongated and sickle shaped (Fig. 1E); lateral sclerite of aedeagus finger-like with basal
thorn-like process (Fig. 1G: s); lingula usually present (as on Fig. 1E: 1).

Intraspecific variability. In most of the taxa in the Merodon serrulatus species
group, the length of pile on the metafemur and the presence of microtrichia on the
scutum and terga is highly variable among specimens of the same species.

Revision of the Merodon serrulatus group 91

i

Figure 5. Merodon serrulatus head. A lateral view, male B dorsal view, male C lateral view, female D dor-
sal view, female. Scale bar: 2 mm.

The Merodon serrulatus species group consists of 13 species, namely M. bequaerti,
M. defectus sp. nov., M. disjunctus sp. nov., M. hirsutus, M. kawamurae, M. medium sp.
nov., VM. nigrocapillatus sp. nov., M. nigropunctum sp. nov., M. opacus sp. nov., M. sacki,
M. serrulatus, M. sophron Hurkmans, 1993, and M. trianguloculus sp. nov.

Taxonomy and nomenclature of the species belonging to the Merodon serrulatus
species group

Merodon bequaerti Hurkmans, 1993
Figs 8A, B, 9A—C, J, 10A, B, 11A—C

Diagnosis. Large (8—11.9 mm), dark brown species with pairs of narrow microtrichose
fasciae on terga 2—4 in males, in some specimens absent; metafemur with long pile on

92 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 6. Male genitalia, surstylus, lateral view. A Merodon serrulatus, Spain B Merodon serrulatus, France
C Merodon serrulatus, Greece (Pindos) D Merodon serrulatus, Greece (Olympos) E Merodon serrulatus,
Greece (Peloponnese) F Merodon serrulatus, Montenegro G Merodon serrulatus, Russia H Merodon me-
dium sp. nov. G, H margin of anterior surstyle lobe marked with arrow. Abbreviation: bp—basolateral

protrusion. Scale bar: 0.2 mm.

ventral margin; the longest pile as long as one third to half of width of metafemur (Fig.
8); apical part of anterior surstyle lobe rhomboid shape, covered with dense, short pile,
and strong dark brown marginal pile on posterior surstyle lobe (Fig. 9A: al, J); females
with very narrow microtrichose fasciae on terga 2—4 and sparse pilosity on ventral
margin of metafemur, only with few longer pile. Similar to Merodon sacki but differs
in a less curved metafemur and generally shorter body pile in males, clearly visible on
tergum 4 (Fig. 10), and by well separated anterior and posterior surstyle lobe (Fig. 9A),
almost fused in M. sacki (Fig. 9D). Related to M. sophron, but differs in more incrassate
metafemur (Fig. 8A, D), longer pile on ventral margin of metafemur in both sexes (Fig.

Revision of the Merodon serrulatus group

ie )

aan eS \ ° peo Se _ ie Uy ad f Jisy > fi: AEST S Gi
OM. bequaerti @ M. defectus sp.nov. @M. disjunctus sp. nov. @M. hirsutus OM. kawamurae @M. medium sp. nov.

O M. nigrocapillatus sp.nov. @M.nigropunctum sp.nov. @M.opacussp.nov. @M. trianguloculus sp. nov.

@ M. sacki O M. serrulatus @ MM. sophron

Figure 7. Distribution map of Merodon serrulatus species group.

ae

Figure 8. Metaleg, lateral view. A Merodon bequaerti, male B Merodon bequaerti, female C Merodon
sacki, male D Merodon sophron, male. Scale bar: 2 mm.

94 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

b

Figure 9. Male genitalia. A Merodon bequaerti, epandrium, lateral view B Merodon bequaerti, epan-

drium, ventral view C Merodon bequaerti, hypandrium, lateral view D Merodon sacki, epandrium, lateral
view E Merodon sacki, epandrium, ventral view F Merodon sacki, hypandrium, lateral view G Merodon
sophron, epandrium, lateral view H Merodon sophron, epandrium, ventral view | Merodon sophron, hypan-
drium, lateral view J Merodon bequaerti, anterior surstyle lobe, lateral view K Merodon sophron, anterior
surstyle lobe, lateral view. Abbreviations: al—anterior surstyle lobe, bp—basolateral protrusion, |—lingula,

pl-posterior surstyle lobe. Scale bar: 0.2 mm.

Revision of the Merodon serrulatus group 95

8), and presence of dense, dark brown marginal pile on apical part of anterior surstyle
lobe (Fig. 9A, J), less dense and light yellow in M. sophron (Fig. 9G, K ).

Redescription (based on the type material and additional specimens). Male. Head.
Antennae black to dark brown; basoflagellomere 1.7—2.1 times as long as wide, and 2.3
times as long as pedicel, concave dorsally with acute apex; fossette dorsolateral (Fig.
11); arista dark and thickened at basal one third, covered with dense microtrichia; arista
1.4—1.7 times as long as basoflagellomere (Fig. 11A, B); face and frons black with gray
microtrichia, face covered with dense whitish gray, and frons with yellowish gray pile;
oral margin shiny with microtrichose lateral areas; lunule shiny black, bare; eye con-
tiguity 10—12 facets long; vertex isosceles, shiny covered with golden microtrichia in
front of ocellar triangle; vertex with long, pale whitish yellow pile mixed with black pile
on the ocellar triangle; ocellar triangle equilateral; occiput shiny, with gray-yellow pile,
covered with a dense, gray microtrichia in ventral half; eyes covered with dense pile.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile; scutum at wing basis with short black pile, in some specimens with fas-
cia of black pile between wing basis; scutum with two or more microtrichose vit-
tae, anteriorly connected and posteriorly reaching the scutellum; posterodorsal part
of anterior anepisternum, posterior anepisternum (except anteroventral angle), ante-
rior anepimeron, dorsomedial anepimeron, and posterodorsal and anteroventral parts
of katepisternum with long, pale yellow pile and grayish microtrichia; wings entirely
covered with microtrichia; wing veins brown; calypteres and halteres pale yellow; legs
without spinae or other protuberances; legs mostly black, except brown tarsi ventrally
in some specimens; pile on legs pale yellow, except black pile at apical one fourth of
metafemur; metafemur curved and incrassate, ca. three times longer than wide; pile on
postero- and anteroventral surface long, and ca. one third to half of width of metate-
mur, slightly longer than pile on dorsal margin (Fig. 8A).

Abdomen. Wide, tapering, 1.2 times longer than mesonotum; terga dark brown to
black, with or without pairs of narrow microtrichose fasciae; tergum 2 with orange lateral
maculae; pile on terga all yellow, except few black pile on medial part of terga 3 and 4 in
some specimens (Fig. 10A, B); sterna dark brown, covered with long whitish/yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, 1.5 times
longer than wide, covered with dense, short pile, and strong dark brown marginal pile
(Fig. 9A: al, J); posterior surstyle lobe oval (Fig. 9A: pl) with basolateral protrusion
(lateral hump) (Fig. 9A, B: bp); hypandrium sickle-shaped, without lateral projections;
lingula small (Fig. 9C: I).

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: antennae with rounded tip, basoflagellomere 1.7—1.9 times
longer than wide (Fig. 11C); frons with microtrichose vittae along eye margins; frons
covered with pilosity of variable color, from mostly gray-yellow until predominantly
black; ocellar triangle covered with black pile; ventral margin of metafemur with sparse
pilosity, only few pile longer (Fig. 8B); lateral side of terga, anterior two third of ter-
gum 2 and all tergum 5 with yellow pile; terga 2-4 with short adpressed black pile and
with very narrow microtrichose fascia.

96 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Figure 10. Abdomen of males. A Merodon bequaerti, dorsal view B Merodon bequaerti, lateral view C
Merodon sacki, dorsal view D Merodon sacki, \ateral view. Scale bar: 2 mm.

Distribution. Merodon bequaerti is distributed in north-western Africa (Algeria,
Libya, Morocco, and Tunisia) (Fig. 7).

Ecology. Preferred environment: unimproved montane grassland, including open,
grassy areas in pine forest or Mediterranean scrub. Flowers visited: no data. Flight pe-
riod: February-June.

Type material. Holotype [original designation by Hurkmans (1993: 194)]: male. Orig-
inal label: “Merodon bequaerti | spec. nov. HOLOTYPE | 3. W Hurkmans 1988.” [red label
handwritten], “Merodon / parietum | Mg 3” [label handwritten], “Noiseux Oran / Algeria /
Dr. J. Bequaert” “23-IV-10” [handwritten on the back side] (MNHN) (See Supplementary
file 1: Figure 1A). Paratype: female. Same label data as holotype (MNHN) (studied).

Revision of the Merodon serrulatus group 97

Figure | 1. Antenna. A Merodon bequaerti, outer side, lateral view, male B Merodon bequaerti, inner side,
lateral view, male C Merodon bequaerti, outer side, lateral view, female D Merodon sacki, outer side, lateral
view, male E Merodon sacki, inner side, lateral view, male F Merodon sacki, dorsal view, male G Merodon
sophron, outer side, lateral view, male H Merodon sophron, inner side, lateral view, male | Merodon sophron,

outer side, lateral view, female. Scale bar: 1 mm.

Other material. ALcErta * 1 @; Kabylie, Tikjda; 36°27'00"N, 4°07'60"E; 28 Jun.
1954; NBCN « 1 <Q; Jijel, Oued el Kebir; 36°35'22"N, 6°16'16"E; 20 May 1981;
NBCN °* 1 @; Jijel, Foce Oued El Kebir; 36°35'45"N, 6°15'29"E; 20 May 1981; I.
Aslan leg.; NBCN 05636 ¢ 1 9; El Kseur, Akfadou; 36°37'60"N, 4°36'00"E; 22-23
May 1981; NBCN 04079.

98 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Lipya * 1 Q; Tripolitania, Garian; 32°10'46"N, 13°01'53"E; “2.500 feet” [760 m
a.s.l.]; 22 Feb. 1954; K. M. Guichard leg.; NHMUK 04353.

Morocco * 1 33; Moyen Atlas, Azrou; 33°25'48"N, 5°12'36"W; 15 Jun. 1928;
R. Benoist leg.; MNHN 22623 * 8 3; Mountain de Beni-Snassen 2; 34°48'43"N,
2°24'08"W; 29 Apr. 2013; A. Vujié, S. Radenkovi¢ leg.; FSUNS D13, D14, D16 to D21.

Tunisia * 1 3; Jundubah, 40 km W from Jendouba; 36°31'54"N, 8°28'25"E; 17
May 1988; ZMUC 02497 © 1 @; same data as for preceding; 36°34'33"N, 9°02'12"E;
ZMUC 02498.

Merodon defectus Vuji¢, Likov & Radenkovi€ sp. nov.
http://zoobank.org/7EDC43D 1-8B17-46D0-8C28-B 18485482741
Figs 12J, K, 13E, 14A, B, D, 15B, 16C

Diagnosis. Medium sized (7.6—10.9 mm), dark species with olive-brown reflection;
antennae dark brown; legs mostly black; basoflagellomere elongated (in males 1.8
times as long as wide) obviously concave dorsally; arista short, 1.6—1.8 times as long
as basoflagellomere (Fig. 12J, K); terga dark brown to black, except pale yellow-orange
lateral maculae on tergum 2; metafemur incrassate, ventrally covered with pilosity of
medium length (Fig. 13E); male genitalia: posterior surstyle lobe with very small lateral
hump (Fig. 14A, B: bp); apical part of anterior surstyle lobe triangular (Fig. 14A: al);
lingula medium sized (Fig. 14D: 1). Similar to Merodon serrulatus from which differs
in reduced lateral hump on posterior surstyle lobe (Fig. 14A, B: bp) (in MZ. serrulatus
distinct, Fig. LA, B: bp, 14C: bp). Morphologically related to M. opacus sp. nov. and
M. hirsutus from which can be distinguished by the presence of yellow-orange lateral
maculae on tergum 2 (in MZ. opacus sp. nov. and M. hirsutus tergum 2 dark). Addition-
ally, differing from M. hirsutus by shorter dorsolateral pile on metafemur (Fig. 13E)
and posterior surstyle lobe with very small lateral hump (Fig. 14A, B: bp), well devel-
oped in M. hirsutus (Fig. 14E, F: bp) and M. opacus sp. nov. (14H, I: bp).

Description. Male. Head. Antennae black to dark brown; basoflagellomere elon-
gated, 1.8 times as long as wide, and 2.3 times as long as pedicel, concave dorsally with
acute apex; dorsolateral fossette large; arista dark brown and thickened at basal one
third, covered with dense microtrichia, 1.6—1.8 times as long as basoflagellomere (Fig.
12J); face black with gray microtrichia, covered with whitish pile; oral margin micro-
trichose, with small, shiny, lateral bare area; lunule shiny black; frons microtrichose,
with yellowish gray pile; eye contiguity ca. eight facets long; vertex isosceles, with long,
pale whitish yellow pile, mixed with black pile on the ocellar triangle; ocellar triangle
isosceles (Fig. 16C); occiput shiny, with gray-yellow pile; eyes covered with dense pile;
vertical triangle: eye contiguity: frons = 3: 1: 3.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile; scutum at wing basis with short black pile and some black pile between
wing basis, from few ones to fascia of black pilosity in some specimens; scutum usually
with two or more microtrichose vittae, anteriorly connected and posteriorly reaching

Revision of the Merodon sertulatus group 99
A B C dy
= Os
D E F i
3,
G H I
N Sy

Ss ae ds ae,

J

Figure 12. Antenna. A Merodon opacus sp. nov., outer side, lateral view, male B Merodon opacus sp. nov.,
inner side, lateral view, male C Merodon opacus sp. nov., dorsal view, male D Merodon opacus sp. nov.,
outer side, lateral view, female E Merodon opacus sp. nov., inner side, lateral view, female F Merodon opacus
sp. nov., dorsal view, female G Merodon disjunctus sp. nov., outer side, lateral view, male H Merodon dis-
junctus sp. nov., inner side, lateral view, male | Merodon disjunctus sp. nov., outer side, lateral view, female
J Merodon defectus sp. nov., outer side, lateral view, male K Merodon defectus sp. nov., outer side, lateral

view, female. Scale bar: 1 mm.

the scutellum; scutum dull; posterodorsal part of anterior anepisternum, posterior an-
episternum (except anteroventral angle), anterior anepimeron, dorsomedial anepimer-
on, and posterodorsal and anteroventral parts of katepisternum with long, pale yellow
pile and grayish microtrichia; wings entirely covered with microtrichia; wing veins
brown; calypteres yellowish; halteres brown-yellow; legs mostly black, except brown
tarsi ventrally in some specimens; pile on legs gray-yellow; metafemur moderately in-

Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 13. Metaleg, lateral view. A Merodon opacus sp. nov., male B Merodon opacus sp. nov., female

C Merodon hirsutus, male D Merodon hirsutus, female E Merodon defectus sp. nov., male. Scale bar: 2 mm.

Revision of the Merodon serrulatus group 101

crassate, ca. three times longer than wide; pile on postero- and anteroventral surface
of medium length; pile on dorsolateral surface dense and length ca. one third to one
fourth of width of metafemur (Fig. 13E).

Abdomen. Tapering, 1.2 times longer than mesonotum; terga dark brown to black,
except pale yellow-orange lateral maculae on tergum 2; terga 2—4 each with a pair of
white microtrichose, oblique fasciae (on tergum 2 triangular); pile on terga long, yel-
low, except some black pile on terga 3 and 4 medially; sterna dark brown, covered with
long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe triangular, ca. two times longer
than wide, covered with dense, short pile (Fig. 14A: al); posterior surstyle lobe with
very small basolateral protrusion (Fig. 14A, B: bp); hypandrium sickle-shaped, without
lateral projections; lingula medium size (Fig. 14D: 1).

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: basoflagellomere ca. two times longer than wide (Fig. 12K),
fossette dorsal; frons with broad microtrichose vittae along eye margins; frons covered
with pilosity of variable color, from mostly gray-yellow to predominantly black; ocellar
triangle covered with black pile; terga with whitish pile, except terga 2-5 medially with
short black pile; microtrichose fasciae on terga 3 and 4 conspicuous (Fig. 15C).

Etymology. Latin adjective defectus (reduced in size, smaller) refers to small baso-
lateral protrusion (lateral hump) on posterior surstyle lobe.

Distribution. Merodon defectus sp. nov. has been identified in western Turkey (Fig. 7).

Ecology. Preferred environment: forest/open ground; thermophilous and evergreen
Quercus forest; Castanea forest, dry Pinus forest; unimproved grassland and tracksides;
coniferous forest with some yellow flowers along a stream [Reemer and Smit (2007)
refer to this last observation as being Merodon alexeji Paramonov, 1925]. Flowers visited:
Ornithogalum spp., Potentilla spp., and Thymus spp. Flight period: May-July.

Type material. Holotype. Turkey * 3; Bozda% mountain, Near Bozdag;
38°22'28"N, 28°04'38"E; 1140 ma.s.L; 7 Jun. 2014; A. Vuji¢, J. A¢anski leg.; FSUNS
06950. Original label: “HOLOTYPE of Merodon | defectus Vuji¢, Likov et / Radenkovi¢
sp.n. 2019” [red label], “Turkey, Bozdag Mountain, / near Bozdag 7/6/2014 /
38.374523 28.077339 1140m / Leg. Vuji¢, Acanski”, “AU305”, “06950” (See Sup-
plementary file 2: Figure 2A). Paratypes. Turkey * 1 3; Muéla, 14 km NE from Aégla,
Lake Kartar; 37°01'50"N, 28°45'09"E; 1600 m a.s.L.; 31 May 2000; J. T. Smit leg.; J.
T. S. coll. 04066 [published in Reemer and Smit (2007) under name Merodon alexeji]
¢ 1 9; Mugla, 14 km NE from Aégla, Lake Kartar; 37°01'50"N, 28°45'09"E; 1600
m a.s.l.; 31 May 2000; J. T. Smit leg.; J. T. S. coll. 04067 [published in Reemer and
Smit (2007) under name Merodon alexeji] * 4 99; Isparta, Yenisarbademli, Melikler
Yaylast 2337°41-389N,.31717 56°E1770 mas lise? Jun. 2006; .R. HayatwA. Vanjic;
O. Demirdézer, J. Acanski leg.; EMIT 12301 to 12304 ® 1 3; Babadag, Near Denizli
valley I; 37°41'43"N, 28°59'35"E; 1870 m a.s.L.; 5 Jul. 2015; A. Vuji¢, S. Radenkovié,
J. Acanski, S. Gékhan, N. Velickovié leg.; FSUNS 09774 * 2 99; Babadag, Near
Denizli valley I; 37°41'43"N, 28°59'35"E; 1870 m a.s.l.; 5 Jul. 2015; A. Vuji¢, S.
Radenkovi¢, J. A¢anski, S. Gdkhan, N. Veli¢kovi¢ leg.; FSUNS 09773, 09775 © 3 3A;

102 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 14. Male genitalia. A Merodon defectus sp. nov., epandrium, lateral view B Merodon defectus sp.
noy., epandrium, ventral view C Merodon serrulatus, epandrium, ventral view D Merodon defectus sp. nov.,
hypandrium, lateral view E Merodon hirsutus, epandrium, lateral view F Merodon hirsutus, epandrium,
ventral view G Merodon hirsutus, hypandrium, lateral view, H Merodon opacus sp. nov., epandrium, lateral
view | Merodon opacus sp. nov., epandrium, ventral view J Merodon opacus sp. nov., hypandrium, lateral
view. Abbreviations: al—anterior surstyle lobe, bp—basolateral protrusion, c—cercus, |-lingula, pl—posterior
surstyle lobe. Scale bar: 0.2 mm.

Isparta, Yenisarbademli, Melikler Yaylasi; 37°41'52"N, 31°17'39"E; 1730 m a.s.l.; 30
Jun. 2015; A. Vuji¢, R. Hayat, O. Dermirézer, A. Uzal leg.; EMIT 09953 to 09955 ¢ 3
Q Q; Isparta, Yenisarbademli, Melikler Yaylasi; 37°41'52"N, 31°17'39"E; 1730 ma.s.L;
30 Jun. 2015; A. Vuji¢, R. Hayat, O. Demirézer, A. Uzal leg.; EMIT 09952, 09956,
09957 © 8 OC; Isparta, Yenisarbademli, Melikler Yaylasi 1; 37°41'52"N, 31°17'39"E;
1730 ma.s.l.; 21 Jun. 2016; R. Hayat, A. Vuji¢, O. Demirézer, J. Acanski leg.; EMIT
12249 to 12251, 12256, 12257, 12260, 12264 * 9 99; Isparta, Yenisarbademli, Me-
likler Yaylasv 13-37°4152"N,-31°17'39" Ey 1730. mass 2) Jun’ 2016; Re-Hayat, A.
Vuji¢, O. Demirézer, J. Acanski leg.; EMIT12253: to.12255, 12258)°12259} 12261 to

Revision of the Merodon serrulatus group 103

Figure 15. Abdomen, dorsal view. A Merodon disjunctus sp. nov., male B Merodon defectus sp. nov.,
female. Scale bar: 2 mm.

12263, 12265 * 3 QQ; Babadag, Near Denizli on the top; 37°42'33"N, 28°59'23"E;
2060 m a.s.l.; 5 Jul. 2015; A. Vujic, S. Radenkovi¢, J. Acanski, S$. Gokhan, N.
Velickovié leg.; FSUNS 09769, 09770, 09772 * 1 2; Bozdag mountain, Near Bozdag;
38°19'58"N, 28°06'35"E; 1570 ma.s.L; 7 Jun. 2014; A. Vuji¢, J. A¢anski leg.; FSUNS
06931 ° 8 od; Bozdag mountain, Near Bozdag; 38°20'50"N, 28°04'08"E; 1170 m
a.s.l.; 7 Jun. 2014; A. Vuji¢, J. Acanski leg.; FSUNS 06952, 06953, 06956, 06957,
06959 to 06962 * 6 99; Bozdag mountain, Near Bozdag; 38°20'50"N 28°04'08"E;
1170 ma.s.L; 7 Jun. 2014; A. Vuji¢, J. Acanski leg.; FSUNS 06954, 06955, 06958,
06963 to 06965 © 1 3; Balikesir, Edremit-Akcay; 39°40'40"N, 26°54'09"E; 27 Jul.
2015; J. Devalez leg.; MAegean 10131.

Merodon disjunctus Vuji¢, Likov & Radenkovi¢ sp. nov.
http://zoobank.org/2A04080B-F1B9-4CE1-A2B1-42AECB902F65
Figs 12G—I, 15A, 16A, B, 17A—C, 18A—C

Diagnosis. Medium sized (8.5—10.8 mm), dark, olive-brown species, covered with
pale yellow pile; males with dichoptic eyes, separated by a distance of 3—5 facets (Fig.
16B); terga 2—4 with pairs of white microtrichose fasciae, differently developed (from
conspicuous to vague) (Fig. 15A); in male basoflagellomere short, 1.2 times longer
than wide, with large fossette extending to the apex of basoflagellomere (Fig. 12G-—I).

Description. Male. Head. Antennae black; basoflagellomere short, 1.2 times as
long as wide, and ca. 1.7 times as long as pedicel, and with rounded apex; large fossette

104 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Figure 16. Head of male. A Merodon disjunctus sp. nov., lateral view B Merodon disjunctus sp. nov.,

dorsal view C Merodon defectus sp. nov., dorsal view. Scale bar: 2 mm.

Figure 17. Merodon disjunctus sp. nov. A thorax, dorsal view, male B metaleg, lateral view, male C met-

aleg, lateral view, female. Scale bar: 2 mm.

dorsomedial and dorsolateral including apex of basoflagellomere (Fig. 12G); arista
black and thickened at basal one third, covered with dense microtrichia, 1.5 times as
long as basoflagellomere (Fig. 12G—H); face and frons black with gray microtrichia;
face covered with dense whitish gray, while frons with mostly black pile (Fig. 16A); oral
margin shiny, with lateral microtrichose area; lunule shiny black, bare; eye dichoptic,
separated by distance of 3—5 facets (Fig. 16B); vertex isosceles, covered with dark gray
microtrichia and long, black pile; ocellar triangle equilateral; occiput shiny covered
with black pile in upper half, ventrally with gray-yellow pile and dense, gray microtri-
chia; eyes covered with dense pile.

‘Thorax. Scutum and scutellum dull, with bronze luster, covered with dense, erect,
yellowish pile, except posterior half medially with few to many black pile intermixed;
in some specimens scutellum with few black pile; scutum with indistinct microtrichose
vittae (Fig. 17A); posterodorsal part of anterior anepisternum, posterior anepisternum
(except anteroventral angle), anterior anepimeron, dorsomedial anepimeron, and pos-
terodorsal and anteroventral parts of katepisternum with long, dark gray pile and gray-

Revision of the Merodon serrulatus group 105

ish microtrichia; wings entirely covered with microtrichia; wing veins yellow-brown;
calypteres and halteres whitish yellow; legs black, except yellow-brown tarsi, tip of
femora and basal part of tibiae; pile on legs mostly yellowish; metafemur moderate inc-
rassate, ca. five times longer than wide; pile on postero- and anteroventral surface long,
ca. half to two thirds of width of metafemur, as same length as pile dorsally (Fig. 17B).

Abdomen. Tapering, ca. 1.2 times longer than mesonotum; terga dark brown to
black, except for a pair of yellow-orange, triangular, lateral maculae on tergum 2; terga
2—4 with conspicuous or with trace of white microtrichose pair of fasciae (variable
character); pile on terga mostly yellow, except terga 3 and 4 medially with black pile;
sterna dark brown to black, covered with long whitish pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid in shape, ca. 1.5
times longer than wide, covered with dense, short pile (Fig. 18A: al); posterior surstyle
lobe oval with basolateral protrusion (lateral hump) (Fig. 18A, B: bp); hypandrium
sickle-shaped, without lateral projections; lingula medium sized (Fig. 18C: 1).

Female. Similar to the male except for normal sexual dimorphism and for the follow-
ing characteristics: basoflagellomere 1.6—1.8 times longer than wide, fossette large, dorso-
lateral (Fig. 121); frons mostly microtrichose and predominantly covered with black pile;
ocellar triangle covered with black pile; microtrichose fasciae on terga 3 and 4 conspicuous.

Etymology. The name derives from the Latin adjective disjunctus meaning sepa-
rated, disconnected which pertains to the dichoptic eyes in the males.

Distribution. Merodon disjunctus sp. nov. has so far only been recorded in Kyr-
gyzstan and Kazakhstan (Fig. 7).

Ecology. Preferred environment: no data. Flowers visited: no data. Flight period:
May-July.

Type material. Holotype. Kyrcyzstan * 3; Talassky Mt.R, Ara Bijik rav., 13 km
NNE Majdantal Pass; 42°22'00"N, 70°00'00"E; 2700 m a.s.l.; 4 Jul. 1988; Milko leg,;
SZMN 05847. Original label: “HOLOTYPE of Merodon | disjunctus Vuji¢, Likov et /
Radenkovi¢ sp.n. 2019” [red label], “NW KIRG., Talassky Mt.R, / Ara Bijik rav., 13
km NNE / Majdantal Pass ~2700 m / 42°22’N 70°57E / 04.07.1998 D. Milko leg.”,
“05847” (See Supplementary file 2: Figure 2B). Paratypes. KazAKHSTAN * 1 4; Ketmen,
Mt. Kirgyzsay; 43°16'60"N, 79°31'00"E; 2200 m a.s.l.; 3 Jun. 2001; M. Hauser leg;
M. H. coll. 02470 © 2 44; Ketmen, Mt. Kirgyzsay; 43°16'60"N, 79°31'00"E; 1800 m
a.s.l.; 1-3 Jun. 2001; M. Hauser leg.; M. H. coll. 02467, 02472 ¢ 1 9; Almaty, Charyn;
43°46'58"N, 79°23'24"E; 20 May 2003; A. Selin leg.; S. K. coll. 03971 ¢ 2 2 9; Kyzyltchy;
46°03'31"N, 80°43'10"E; 21 May 2004; A. Selin leg.; S. K. coll. 03970, 02460 ¢ 1 4;
Kyzyltchy; 46°03'31"N, 80°43'10"E; 21 May 2004; A. Selin leg.; S. K. coll. 02461 1 4;
Talasskip hr., R. Kara-Bura; 1600 m a.s.l.; 18 Jul. 1968; Pek leg.; SZMN 05815.

Kyreyzstan * 1 6; Tchatkal Valley, 4 SW Ajgyr-Dzhal vill; 41°42'00"N,
70°57'00"E; 1800 m a.s.l.; 12 Jul. 1998; Milko leg.; SZMN 05848 1 GO; Issyk Kul,
Chong Kemin-Tal; 42°40'60"N, 75°55'00"E; 1350 ma.s.1.; 3 Jun. 1998; M. Kraus M.
leg.; NBCN 02468 ¢ 1 Q; Issyk Kul, Chong Kemin-Tal; 42°42'00"N, 75°54'00"E; 1350
ma.s.l.; 3 Jun. 1998; M. Kraus leg.; NBCN 02464 ¢ 1 G; Issyk Kul, Chong Kemin-Tal;
42°42'00"N, 75°54'00"E; 1350 ma.s.l.; 3 Jun. 1998; M. Kraus leg.; NBCN 02469.

106 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Figure 18. Male genitalia. A Merodon disjunctus sp. nov., epandrium, lateral view B Merodon disjunctus
sp. noy., epandrium, ventral view C Merodon disjunctus sp. nov., hypandrium, lateral view D Merodon
kawamurae, epandrium, lateral view E Merodon kawamurae, epandrium, ventral view F Merodon kawa-
murae, hypandrium, lateral view. Abbreviations: al—anterior surstyle lobe, bp—basolateral protrusion, |-

lingula, pl—posterior surstyle lobe, s—lateral sclerite of aedeagus. Scale bar: 0.2 mm.

Merodon hirsutus Sack, 1913
Figs 13C, D, 14E-G, 19A—D, 20A, B, 21E-G

Diagnosis. Medium sized (8.1—10.4 mm), dark species with olive-brown reflection;
antennae dark brown; legs mostly black; basoflagellomere elongated (ca. two times as
long as wide) obviously concave dorsally; arista 1.6—1.7 times as long as basoflagel-
lomere (Fig. 19A—D); terga dark brown to black; metafemur incrassate, covered with
short pilosity ventrally, and with long pile on dorsolateral surface (Fig. 13C); male
genitalia: posterior surstyle lobe with lateral hump (Fig. 14E, F: bp); apical part of
anterior surstyle lobe rhomboid (Fig. 14E: al); lingula large (Fig. 14G: 1). Similar to
Merodon serrulatus from which differs in dark tergum 2 (in M. serrulatus with small
pale lateral maculae, at least in females). Morphologically related to M. opacus sp. nov.
from which can be distinguished by longer dorsolateral and ventral pile on metafemur
(Fig. 13C) and longer pile on terga (Fig. 21, G); females with mostly shiny frons (Fig.
20B) (in M. opacus sp. nov. dull, covered with dense microtrichia).

Redescription (based on lectotype and additional specimens from the type local-
ity, Syria, and Israel). Male. Head. Antennae black to dark brown; basoflagellomere
elongated ca. two times as long as wide, and 2.2—2.5 times as long as pedicel, concave

Revision of the Merodon serrulatus group 107

dorsally with acute apex; large fossette dorsolateral and dorsomedial; arista dark and
thickened at basal one third, covered with dense microtrichia, 1.6—1.7 times as long as
basoflagellomere (Fig. 19A—C); face black with gray microtrichia, covered with whit-
ish pile; frons mostly shiny, with yellowish gray pile; oral margin microtrichose, with
small, shiny lateral area; lunule shiny black, bare; eye contiguity 12-14 facets long;
vertex isosceles, with long, pale whitish yellow pile, mixed with black pile on the ocellar
triangle; ocellar triangle equilateral; occiput with gray-yellow pile; eyes covered with
dense pile (Fig. 20A); vertical triangle: eye contiguity: frons = 1.4:1:2.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile; scutum at wing basis with short black pile; scutum usually with two or
more microtrichose vittae, anteriorly connected and posteriorly reaching the scutel-
lum; scutum dull; posterodorsal part of anterior anepisternum, posterior anepister-
num (except anteroventral angle), anterior anepimeron, dorsomedial anepimeron, and
posterodorsal and anteroventral parts of katepisternum with long, pale yellow pile and
grayish microtrichia; wings entirely covered with microtrichia; wing veins brown; ca-
lypteres and halteres yellowish; legs mostly black, except brown tarsi ventrally in some
specimens; pile on legs pale yellow, except black pile at apical one third of metafemur;
metafemur moderately incrassate, ca. three times longer than wide; pile on postero-
and anteroventral surface very short; pile on dorsolateral surface long and dense ca. as
half of width of metafemur (Fig. 13C).

Abdomen. Tapering, ca. 1.2 times longer than mesonotum; terga dark; terga 2—4
each with a pair of white microtrichose, wide, oblique fasciae (on tergum 2 triangular);
pile on terga long, all yellow (Fig. 21F); sterna dark brown, covered with long whitish
yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, ca. two
times longer than wide, covered with short pile (Fig. 14E: al); posterior surstyle lobe
oval with basolateral protrusion (lateral hump) (Fig. 14E, F: bp); hypandrium sickle-
shaped, without lateral projections; lingula large (Fig. 14G: 1).

Female. Similar to the male except for normal sexual dimorphism and for the fol-
lowing characteristics: antennae with rounded tip, basoflagellomere 1.8—2 times longer
than wide, fossette dorsolateral (Fig. 19D); frons with narrow microtrichose vittae
along eye margins; frons covered with variable pilosity, from mostly gray-yellow to
predominantly black; ocellar triangle covered with black pile; metafemur with shorter
pile on dorsolateral surface (Fig. 13D); terga mostly with yellowish pile on tergum 2
and with whitish pile on terga 3—5, except terga 2—4 medially with short black pile;
microtrichose fasciae on terga 3 and 4 narrower (Fig. 21F, G).

Distribution. Merodon hirsutus is distributed in Israel, Syria, and south-eastern
Turkey (Fig. 7).

Ecology. Preferred environment: no data. Flowers visited: no data. Flight period:
March-June.

Type material. Described by Sack (1913: 435) based on unspecified number of
specimens. Lectotype [designated by Hurkmans (1993)]: male, “Jebel al Aqra [35° 35’
N, 36° 15° E] vi.85 [1885], N. Syria, Dr. E. Leuthner / Lampetia hirsuta Sack det.
Sack” (NHMW) (studied).

108 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

A B

Figure 19. Antenna. A Merodon hirsutus, outer side, lateral view, male B Merodon hirsutus, inner side,
lateral view, male C Merodon hirsutus, dorsal view, male D Merodon hirsutus, outer side, lateral view, fe-
male E Merodon kawamurae, outer side, lateral view, male F Merodon kawamurae, inner side, lateral view,
male G Merodon kawamurae, dorsal view, male H Merodon kawamurae, outer side, lateral view, female
| Merodon medium sp. nov., outer side, lateral view, male J Merodon medium sp. nov., outer side, lateral
view, female. Scale bar: 1 mm.

Figure 20. Head, dorsal view. A Merodon hirsutus, male B Merodon hirsutus, female C Merodon opacus
sp. nov., male. Scale bar: 2 mm.

Revision of the Merodon serrulatus group

Figure 21. Abdomen. A Merodon opacus sp. nov., dorsal view, male B Merodon opacus sp. nov., dorsal
view, female C Merodon opacus sp. nov., lateral view, male D Merodon opacus sp. nov., lateral view, female
E Merodon hirsutus, lateral view, male F Merodon hirsutus, dorsal view, female G Merodon hirsutus, \ateral

view, female. Scale bar: 2 mm.

110 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Other material. Israzt ¢ 1 9; Hefa, “Ma‘yan Zevi” [Ma Yan Zevi]; 32°34'00"N,
34°55'60"E; 17 Apr. 1980; TAUI 04162.

Syria * 1 4; Jebel al Aqra; 35°55'18"N, 35°57'51"E; Jun. 1985; D. E Leuthner
leg.; NHMW 02479.

Turkey * 1 3 Icel, Icel-Tasucu, Silifke; 36°22'17"N, 33°54'54"E; 300 m as.L;
17 Mar. 1984; FSUNS 04161 ¢ 1 9; Erdemli; 36°44'59"N, 34°11'51"E; 8 Jun. 2008;
Skorpik leg.; MNHN 17917 © 3 29; Pozanti-Tekir; 37°31'05"N, 34°47'42"E; 6 Jun.
2008; M. Kafka leg.; M. B. coll. 17918 to 17920 © 1 9; Kahramanmaras, Andirin,
Beyolugu village; 37°45'00"N, 36°17'00"E; 1400 m a.s.L; 7 Jun. 2002; S. Saribryik
leg.; S. S. coll. 02482 © 3 $4; same data as for preceding; S. S. coll. 17910 to 17912
¢ 2 29; Kahramanmaras, Andirin, Cigsar village; 37°45'00"N 36°18'00"E; 1400 m
a.s.l.; 7 Jun. 2002; S. Saribiyik leg.; S. S. coll. 17914, 17916 © 2 4; same data as for
preceding: S: 5. coll-.179135-17915:

Merodon kawamurae Matsumura, 1916
Figs 18D—-F, 19E-H, 224A, B, 23E, F

Lampetia micromegas Hervé-Basin, 1929: 111 —syn. published by Hurkmans 1993: 165.

Diagnosis. Medium sized (7.7—11.2 mm), with olive-brown reflection; antennae red-
dish brown; body pile predominantly pale, except some black pile on vertex and terga
2-4 medially; basoflagellomere short, ca. 1.2 times as long as wide, with large dorsal
to dorsolateral fossette, and short arista (Fig. 19E-H); tergum 2 with reddish yellow
lateral maculae; tergum 3 laterally reddish or brown; metafemur incrassate with long
pilosity as long as half of width of metafemur in male and as one third of width of
metafemur in female (Fig. 22A, B); male genitalia: posterior surstyle lobe with small
lateral hump (Fig. 18E: bp); apical part of anterior surstyle lobe rhomboid (Fig. 18D:
al); lingula large (Fig. 18F: I), lateral sclerite of aedeagus elongated (Fig. 18F: s).

Redescription (based on the types of Merodon micromegas and additional material
from China). Male. Head. Antennae reddish brown; basoflagellomere short, ca. 1.2
times as long as wide, and ca. two times as long as pedicel, straight dorsally with acute
apex; dorsal to dorsolateral fossette large; arista reddish brown and thickened at basal
one third, covered with dense microtrichia, ca. 1.3 times as long as basoflagellomere
(Fig. 19E-G); face and frons black with gray microtrichia, face covered with dense
whitish, and frons with yellowish white pile; lunule shiny black, bare; vertex isosceles,
dull, in front of anterior ocellus covered with dense microtrichia; vertex with long, pale
yellow pile, in some specimens mixed with black or dark gray pile on the ocellar tri-
angle; ocellar triangle isosceles; eyes covered with dense pile; occiput with gray-yellow
pile, ventrally covered with a dense, gray microtrichia; eye contiguity ca. ten facets
long; vertical triangle: eye contiguity: frons = 2.5: 1: 2.5.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect
yellow pile; scutum usually with indistinct microtrichose vittae; posterodorsal part of

Revision of the Merodon serrulatus group 111

Figure 22. A, B metaleg, C, D metatrochanter, metafemur and metatibia, lateral view. A Merodon

kawamurae, male B Merodon kawamurae, female C Merodon medium sp. nov., male D Merodon medium
sp. nov., female. Scale bar: 2 mm.

anterior anepisternum, posterior anepisternum (except anteroventral angle), anterior
anepimeron, dorsomedial anepimeron, and posterodorsal and anteroventral parts of
katepisternum with long, dense pale yellow pile and grayish microtrichia; wings en-
tirely covered with microtrichia; wing veins reddish brown; calypteres and halteres pale
yellow; legs mostly black, except tip of femora and basal part of tibiae and brown tarsi
ventrally; pile on legs pale yellow; metafemur incrassate and curved, ca. three times
longer than wide; long pile on postero- and anteroventral surface ca. as half of width of
metafemur, approximately the same length as pile on dorsal surface (Fig. 22A).

Abdomen. Broad, tapering, 1.2 times longer than mesonotum; terga dark, except
for a pair of reddish yellow, triangular, lateral maculae on tergum 2 (and in some speci-
men on 3); terga 2—4 each with a pair of white microtrichose, wide, usually oblique fas-
ciae; pile on terga all yellow, except black pile on tergum 3 medially, and on tergum 2
posteriorly and tergum 4 anteriorly in some specimens (Fig. 23E); sterna dark brown,
covered with long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, covered with
dense, short pile (Fig. 18D: al); posterior surstyle lobe oval with small basolateral pro-
trusion (lateral hump) (Fig. 18E: bp); hypandrium sickle-shaped, without lateral pro-
jections; lingula large (Fig. 18F: 1); lateral sclerite of aedeagus elongated (Fig. 18F: s).

Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Figure 23. Body parts, dorsal view. A Merodon trianguloculus sp. nov., thorax, male B Merodon triangu-
loculus sp. nov., thorax, female C Merodon trianguloculus sp. nov., abdomen, male D Merodon triangulocu-
lus sp. nov., abdomen, female E Merodon kawamurae, abdomen, male F Merodon kawamurae, abdomen,

female. Scale bar: 2 mm.

Revision of the Merodon serrulatus group 113

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: antennae with rounded tip, fossette dorsolateral (Fig. 19D);
frons microtrichose, covered with mostly gray-yellow pile; ocellar triangle covered
with black pile; long pile on postero- and anteroventral surface ca. as half of width of
metafemur (Fig. 22B); microtrichose fasciae on terga 2—4 narrower (Fig. 23F).

Distribution. Merodon kawamurae is known from Japan and China (Fig. 7). This
is the only species of the genus Merodon in eastern Palaearctic.

Ecology. Preferred environment: no data. Flowers visited: no data. Flight period:
April-May.

Type material. Merodon kawamurae was described after an unknown number of
specimens from Kumamoto, Kyushu, Japan, leg. Kawamura. Matsumura’s type mate-
rial is held at the Hokkaido University, Department for Systematic Entomology, at
Sapporo, Japan, but the type material was inaccessible for this study.

Merodon micromegas Lectotype [designated by Hurkmans (1993)]: “Tchen-Kiang,
13.iv.1918 / Lampetia micromegas H. B. type” (MNHN) (studied).

Paralectotypes (Lampetia micromegas). CHINA * 1 3; Chemo; 33°44'32"N,
103°23'45"E; 25 Apr. 1918; MNHN 02520 1 3; Chemo; 33°44'32"N, 103°23'45"E;
26 Apr. 1918; MNHN 02521 * 1 2; Chemo; 33°44'32"N, 103°23'45"E; 23 Apr.
1918; MNHN 02522 ¢ 1 9; Shia-Shu; 10 May 1918; MNHN 02525 ¢ 1 9; Jiangsu,
Nanking; 32°00'27"N, 118°57'22"E; 6 May 1918; MNHN 02524.

Other material. CHINA * 1 9; Ningpo; 29°44'29"N, 121°06'02"E; 29 Apr. 1925;
J. T. Chu leg.; NMNH 05118 ¢ 1 3; Jiangsu, Nanking; 32°00'27"N, 118°57'22"E;
1981; H. Jettmar leg; NHMW 02516 © 1 Q; Jiangsu, Nanking; 32°00'27"N,
118°57'22"E; 15 Apr. 1918; NBCN 02518 © 1 Q; same data as for preceding; 16 Apr.
1918; MNHN ° 16 46; Chenkiang; 32°08'24"N, 119°23'25"E; 1-13 Apr. 1918;
MNHN ° 12 99; same data as for preceding; MNHN * 6 4; Chemo; 33°44'32"N,
103°23'45"E; 23 Apr. 1918; MNHN ¢ 4 99; same data as for preceding; MNHN e
1 2; Chemo; 33°44'32"N, 103°23'45"E; 23 Apr. 1918; NBCN ¢ 1 Q; same data as
for preceding; 26 Apr. 1918; NBCN ¢ 1 3; Hoachan; 16 May 1918; MNHN #2 29;
same data as for preceding; MNHN Ȣ 3 44; Shia-Shu; 22 Apr. 1918; MNHN.

Merodon medium Vuji¢, Likov & Radenkovic sp. nov.
http://zoobank.org/E384B35E-E377-49B6-90CE-DEBDCBCEDDCB
Figs 6H, 19], J, 22C, D, 24A, B, 25A—C

Diagnosis. Large species (10.3-13 mm) with wide dark brown abdomen and yellow-
orange maculae on lateral sides of tergum 2 (Fig. 24A, B); basoflagellomere elongated,
ca. 2.5 times longer than broad (Fig. 19 I, J,); metafemur incrassate (Fig. 22C, D);
terga 2—4 with conspicuous microtrichose fasciae (Fig. 24A, B). Similar to some popu-
lations of Merodon serrulatus, but clearly differs in shape of abdomen: relation between
maximum width of tergum 2 and its medial length is 3.3 in male and 3.5 in female of
M. medium sp. nov. compared with 2.3 in M. serrulatus male and 2.7 in female; male

114 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

genitalia: anterior surstyle lobe with concave margin in M. medium sp. nov. (Fig. 6H:
marked with arrow), convex in M. serrulatus (Fig. 6G: marked with arrow); apical
microtrichose area of anterior surstyle lobe 2.5 times broader than long in M. medium
sp. nov. (Fig. 6G: al), less than one time in M. serrulatus (Fig. 6G: al); molecular data
and distribution (7. medium sp. nov. is an endemic to the island of Crete in Greece).

Description. Male. Head. Antennae black to dark brown; basoflagellomere elon-
gated ca. 2.2 times as long as wide, and ca. 2.5 times as long as pedicel, concave dor-
sally with acute apex; dorsolateral fossette narrow; arista dark and thickened at basal
one third, covered with dense microtrichia, ca. 1.5 times as long as basoflagellomere
(Fig. 191); face and frons black with gray microtrichia, face covered with dense whitish
gray, and frons with yellowish gray pile; oral margin microtrichose with shiny lateral
areas; lunule shiny black, bare; vertex shiny black, except microtrichose area in front
of anterior ocellus; vertex isosceles, with long, pale whitish yellow pile, mixed with few
black pile on the ocellar triangle; ocellar triangle equilateral; eyes covered with dense
pile; occiput with gray-yellow pile, ventrally covered with a dense, gray microtrichia;
eye contiguity 10-12 facets long; vertical triangle: eye contiguity: frons = 1.2: 1: 2.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile, except sides of scutum at wing basis with patch of short black pile and fascia
of black pile between wing basis; scutum with two or more microtrichose vittae, anteri-
orly connected and posteriorly reaching the scutellum; scutum dull; posterodorsal part
of anterior anepisternum, posterior anepisternum (except anteroventral angle), anterior
anepimeron, dorsomedial anepimeron, and posterodorsal and anteroventral parts of
katepisternum with long, pale yellow pile and grayish microtrichia; wings entirely cov-
ered with microtrichia; wing veins brown; calypteres and halteres yellowish; legs mostly
black, except brown tarsi ventrally in some specimens; pile on legs pale yellow, except
few black pile in apical fifth of metafemur in some specimens; metafemur incrassate, ca.
three times longer than wide; pile on postero- and anteroventral surface short, except
few sparse pile approximately the same length as pile on dorsal surface (Fig. 22C).

Abdomen. Broad, tapering, 1.2 times longer than mesonotum; terga dark, except
for a pair of yellow-orange, triangular, lateral maculae on tergum 2; terga 2—4 each with
a pair of white microtrichose, oblique fasciae (on tergum 2 more triangular); pile on
terga all yellow (Fig. 24A); sterna dark brown, covered with long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, 1.5 times
longer than wide, covered with dense, short pile (Fig. 25.A: al) and with concave mar-
gin (Fig. GH: marked with arrow); posterior surstyle lobe oval with basolateral protru-
sion (lateral hump) (Fig. 25B: bp) and basal hook-like extension (Fig. 25A: marked
with arrow); hypandrium sickle-shaped, without lateral projections; lingula medium
sized (Fig. 25C: 1).

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: antennae with rounded tip, basoflagellomere ca. two times
longer than wide, (Fig. 19J); frons with broad microtrichose vittae along eye margins;
frons covered with pilosity of variable color, from mostly gray-yellow until predomi-
nately black pile; ocellar triangle covered with black pile; metafemur incrassate, pile on

Revision of the Merodon serrulatus group 115

Figure 24. Merodon medium sp. nov., abdomen, dorsal view. A male B female. Scale bar: 2 mm.

postero- and anteroventral surface short (Fig. 22D); terga pale yellow pilose at lateral
sides, anterior two thirds of tergum 2 and all terga 4 and 5; terga 2 and 3 medially with
short adpressed black pile; microtrichose fasciae on terga 3 and 4 broad (Fig. 24B).

Etymology. Medium (middle, center) refers to the species’ distribution, being the
only taxon of the group found on Crete, in the middle of Mediterranean Sea.

Distribution. Merodon medium sp. nov. is endemic to the Greek island of Crete
(Fig. 7).

Ecology. Preferred environment: forest/open ground; evergreen oak forest, dry
Pinus forest; scrub with Pistacia lentiscus L.; well-vegetated, unimproved grassland.
Flowers visited: Ornithogalum spp., Potentilla spp. and Thymus spp. Flight period: May.

Type material. Holotype. Greece * @; Crete, Chania, Omalos plain;
35-19 2ileN23° 55150) ES 28-May 2014 AL Mujiceleg sak SWNS<06729-7 Original
label: “HOLOTYPE of Merodon | medium Vuji¢, Likov et / Radenkovié sp.n. 2019”
[red label], “Greece, Crete, Chania, / Omalos plain / 28.05.2014. 35.322593 /
23.930496 Leg. Vuji¢”, “AU298”, “06729” (See Supplementary file 3: Figure 3).
Paratypes. GREECE ® 1 9; Crete, Chania, Imbors; 35°15'08"N, 24°10'28"E; 27 May
2014; A. Vujic leg.; FSUNS 06706 ° 1 4; Crete, Chania, Omalos plain; 35°19'06"N
23°54'51"E; 28 May 2014; A. Vuji¢ leg.; FSUNS 06723 © 1 4; Crete, Chania, Om-
alos plain; 35°19'21"N, 23°55'50"E; 28 May 2014; A. Vuji¢ leg.; FSUNS 06731 ©
1 Q, Crete, Chania, Mescla, 35°24'05"N 23°56'26"E; 28 May 2014; A. Vujié leg.;
FSUNS 06718.

116 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

al oe ‘

Figure 25. Male genitalia. A Merodon medium sp. nov., epandrium, lateral view B Merodon medium
sp. nov., epandrium, ventral view C Merodon medium sp. nov., hypandrium, lateral view D Merodon
nigrocapillatus sp. nov., epandrium, lateral view E Merodon nigrocapillatus sp. nov., epandrium, ventral
view F Merodon nigrocapillatus sp. nov., hypandrium, lateral view. Abbreviations: al—anterior surstyle

lobe, bp—basolateral protrusion, |-lingula; arrow marks the hook-like extension in A. Scale bar: 0.2 mm.

Merodon nigrocapillatus Vuji¢, Likov & Radenkovic¢ sp. nov.
http://zoobank.org/09B368F3-GBCF-4CF 1-95E1-8AC43C14B1D4
Figs 25D-E, 26A—D, 27A, 28C, D

Diagnosis. Medium sized (10.2—10.8 mm), black and shiny species, covered with most-
ly black pile on scutum, terga and legs in both sexes (Fig. 26); antennae dark; legs black;
male dichoptic (Fig. 27A); basoflagellomere short, 1.3 times longer than wide (Fig.
28C); terga black, without pale lateral maculae on tergum 2; microtrichose fasciae on
terga 2—4 very narrow or absent in males (Fig. 26A) and narrow in females (Fig. 26C).
Description. Male. Head. Antennae black; basoflagellomere short, 1.3 times as
long as wide, and ca. two times as long as pedicel, with rounded apex; fossette dorso-
lateral; arista black and thickened at basal one third, covered with dense microtrichia,
ca. two times as long as basoflagellomere (Fig. 28C); face and frons black with gray
microtrichia, face covered with dense whitish gray or mixed black and whitish gray,
and frons mostly with black pile; oral margin shiny, with lateral microtrichose area;
lunule shiny black, bare; vertex isosceles, shiny black, except in front of anterior ocellus
covered with microtrichia; vertex with long, black pile; ocellar triangle equilateral; eyes

Revision of the Merodon serrulatus group Ly?

Figure 26. Merodon nigrocapillatus sp. nov., body parts. A dorsal view, male B lateral view, male C dorsal

view, female D lateral view, female. Scale bar: 2 mm.

covered with dense whitish pile; occiput shiny covered with black pile in upper half,
ventrally with gray-yellow pile and dense, gray microtrichia; eye dichoptic, separated
by distance of three facets (Fig. 27A).

Thorax. Scutum and scutellum black, shiny, covered with dense, erect, black pile;
scutum without microtrichose vittae (Fig. 26A); posterodorsal part of anterior anepis-
ternum, posterior anepisternum (except anteroventral angle), anterior anepimeron,
dorsomedial anepimeron, and posterodorsal and anteroventral parts of katepisternum
with long, dark gray or black pile and grayish microtrichia; wings entirely covered with
microtrichia, except bare area at basal one third; wing veins dark brown; calypteres
gray; halteres blackish; legs black (Fig. 26B); pile on legs mostly black or dark gray;
metafemur curved and medium incrassate, ca. four times longer than wide; pile on
postero- and anteroventral surface ca. one third to half of width of metafemur, slightly
shorter than pile dorsally.

Abdomen. Tapering, 1.2 times longer than mesonotum; terga completely dark;
terga 3 and 4 without, or with indistinct pair of white microtrichose fasciae; pile on
terga mostly black, except anteromedial part of tergum 2 partly covered with whitish
pile (Fig. 26A); sterna dark brown to black, covered with long black and whitish pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, ca. two times
longer than wide, covered with dense, short pile (Fig. 25D: al); posterior surstyle lobe

118 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 27. Head of male, dorsal view. A Merodon nigrocapillatus sp. nov. B Merodon nigropunctum sp.
noy. Scale bar: 2 mm.

oval with basolateral protrusion (lateral hump) (Fig. 25D, E: bp); hypandrium sickle-
shaped, without lateral projections; lingula large (Fig. 25F: 1).

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: basoflagellomere 1.5 times longer than wide, fossette narrow
(Fig. 28D); pile on face of variable color, from black to whitish; frons mostly microtri-
chose, covered with black pile; ocellar triangle covered with black pile; microtrichose
fasciae on terga 2-4 narrow (Fig. 26C); thorax on lateral sides with variable pilosity
color, from black to whitish.

Etymology. The name nigrocapillatus is derived from Latin adjective niger mean-
ing black, dark and Latin noun capillatus meaning long-haired, referring to the long,
black body pile of this species.

Distribution. Merodon nigrocapillatus sp. nov. has only been recorded in Tajikistan
(Fig. 7).

Ecology. Preferred environment: open areas at high altitudes, unimproved grass-
land (Fig. 35D). Flowers visited: white Apiaceae. Flight period: June-July.

Type material. Holotype. Tajikistan * 3 Varzob, Kalon; 39°03'36"N,
68°52'12"E; 2440 m a.s.l.; 1-4 Jul. 2017; A. Barkalov leg.; SZMN 22625. Original
label: “HOLOTYPE of Merodon | nigrocapillatus Vuji¢, Likov / et Radenkovié sp.n.
2019” [red label], “Taszkuucran, Bapso6ckoe / ym., 3 KM c.-B. KuMTAaKa / Kaaon,
2440m u.y.m. 39,06° / N, 68,87° 1-4.07.2017 / C6.A. Bapxaaos”, “2017 / sp. LIA.
Barkalov det., 201” [label partly handwritten], “22625” (See Supplementary file 1:
Figure 1B). Paratypes. Tayrkistan * 1 9; Varzob, Kalon; 39°03'00"N, 68°52'48"E;
2484 ma.s.l.; 7-12 Jul. 2017; A. Barkalov leg.; SZMN 22626 ¢ 1 9; 65km N of Du-
shanbets7side ANZOR pass’59 03: 000N 268? 192" FE: 2380 meats a2 1 Jul. 20105. Je
Dils, J. Faes leg.; G. V. W. coll. 10397 © 1 9; Iskanderkul kishlak, Sarytag; 39°03'00"N,
68°19'12"E; 2374 m asl; 14 Jun. 2018; A. Barkalov leg.; SZMN 24506 ¢ 4 eter
Varzob Canyon, 3 km N-E Kalon kishlak; 39°03'36"N, 68°52'12"E; 2440 m a.s.L.;
A. Barkalov leg.; SZMN ¢ 14 99; same data as for preceding; SZMN ¢ 2 29; same

Revision of the Merodon serrulatus group 119

data as for preceding; 4 and 7 Jul. 2017; V. Zinchenko leg.; SZMN * 3 99; same data
as for preceding; 4 and 7 Jul. 2018; SZMN ¢ 1 9; Varzob Canyon, 3 km N-E Kalon
kishlak; 39°03'36"N, 68°52'12"E; 2356 ma.s.l.; 5 Jul. 2018; A. Barkalov leg.; SZMN
24502 © 1 9; same data as for preceding; 7 Jun. 2018; SZMN 24505 * 1 9; same data
as for preceding; ~2400 ma.s.1.; 28 Jun. 2018; V. Zinchenko leg.; SZMN 24503 © 1 9;
same data as for preceding; 30 Jun. 2018; SZMN 24504 ¢ 1 9; Varzob Canyon, 4 km
N-E Kalon kishlak; 39°04'48"N, 68°51'36"E; 3375 m a.s.l.; A. Barkalov leg.; SZMN
¢ 1 Q; Iskanderkul, Zmeinoe Lake; 39°05'20"N, 68°22'08"E; -2217 ma.s.l.; 16 Jun.
2018; V. Zinchenko leg.; SZMN 24507.

Merodon nigropunctum Vujic, Likov & Radenkovié sp. nov.
http://zoobank.org/13F2CA9B-BD91-4FD6-B66A-B89DE0F 13726
Figs 27B, 28A, B, 29A—C, 30A—C

Diagnosis. Medium sized (10.3 mm), bluish species (Fig. 29A, B) with dark macula
on medial part of wing (Fig. 29C); basoflagellomere narrow and elongated, 1.8 times
as long as wide, rounded at the tip (Fig. 28A, B); arista long, two times as long as baso-
flagellomere; body pile whitish; posterior half of scutum with square shaped area of
black pile medially; terga 2-4 covered with black pile medially, except whitish pilosity
on conspicuous silver microtrichose fasciae (Fig. 29A).

Description. Male. Head. Antennae black to dark brown; basoflagellomere nar-
row and elongated, 1.8 times as long as wide, and 2.5 times as long as pedicel, with
rounded tip; large fossette dorsomedial and dorsolateral (Fig. 28A, B); arista dark and
thickened at basal one third, covered with dense microtrichia; arista long, ca. two
times as long as basoflagellomere (Fig. 28A, B); face and frons black covered with
whitish pile; face covered with indistinct whitish gray microtrichia; frons with dense
whitish microtrichia; lunule shiny black, bare; vertex isosceles, with long whitish pile
and black pilosity on the ocellar triangle; ocellar triangle equilateral; eyes covered with
long, dense, whitish pile (Fig. 27B); occiput with whitish pile, covered with a dense,
silver microtrichia along eye margin; eye contiguity short, approximately five facets
long; vertical triangle: eye contiguity: frons = 4.5: 1: 4.5.

Thorax. Scutum and scutellum black with bluish luster, covered with dense, erect,
white pile including wing basis; posterior half of scutum with square shaped area of
black pile medially; scutum with indistinct microtrichose vittae; posterodorsal part of
anterior anepisternum, posterior anepisternum (except anteroventral angle), anterior
anepimeron, dorsomedial anepimeron, and posterodorsal and anteroventral parts of
katepisternum with long, whitish pile and grayish microtrichia; wings entirely covered
with microtrichia; wing veins dark brown; wing with distinct dark area in apical half
(Fig. 29C); calypteres whitish; halteres yellowish, with darker capitulum; legs mostly
black, except dark brown tarsi ventrally; pile on legs mostly whitish mixed with black
ones on femora; metafemur moderately incrassate, ca. three times longer than wide;
pile on postero- and anteroventral surface very long, and ca. two thirds of width of
metafemur, approximately the same length as pile on dorsal surface.

120 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 28. Antenna. A Merodon nigropunctum sp. nov., outer side lateral view, male B Merodon nigro-
punctum sp. nov., inner side, lateral view, male C Merodon nigrocapillatus sp. nov., outer side, lateral view,

male D Merodon nigrocapillatus sp. nov., outer side, lateral view, female. Scale bar: 1 mm.

Figure 29. Merodon nigropunctum sp. nov., male. A body, dorsal view B body, lateral view C left wing,

dorsal view. Scale bar: 2 mm.

Revision of the Merodon serrulatus group 124

Figure 30. Male genitalia. A Merodon nigropunctum sp. nov., epandrium, lateral view B Merodon nigro-
punctum sp. nov., epandrium, ventral view C Merodon nigropunctum sp. nov., hypandrium, lateral view.

Abbreviations: al—anterior surstyle lobe, bp—basolateral protrusion, |-lingula. Scale bar: 0.2 mm.

Abdomen. Tapering, 1.2 times longer than mesonotum; terga dark brown to black;
terga 2—4 each with a pair of white microtrichose, wide, oblique fasciae (on tergum 2
triangular); pile on terga long, whitish laterally and at microtrichose fasciae, medially
black (Fig. 29A); sterna dark brown, covered with long whitish pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, 1.5 times
longer than wide, covered with dense, short pile (Fig. 30A: al); posterior surstyle lobe
oval with basolateral protrusion (lateral hump) (Fig. 30A, B: bp); hypandrium sickle-
shaped, without lateral projections; lingula large (Fig. 30C: 1).

Female. Unknown.

Etymology. The word nigropunctum is derived from the Latin words niger (black,
dark whitish) and punctum (dot/spot) referring to the dark macula on the wing as an
important diagnostic character of this new species.

Distribution. Merodon nigropunctum sp. nov. was recorded at only one locality in
Uzbekistan (Fig. 7).

Ecology. Preferred environment: no data. Flowers visited: no data. Flight period: May.

Type material. Holotype. UzBEKISTAN * 4; KadamZai, S of Fergana; 40°20'00"N,
71°47'39"E; 21 May 1980; Z. Padr leg.; NMPC 18248. Original label: “HOLO-
TYPE of Merodon | nigropunctum Vujic, Likov / Radenkovié sp.n. 2019” [red label],
“C.ASIA, Uzbekistan / Kadanzai, S of Fergana / 21.5.1980 leg.Z.Padr”, “18248” (See
Supplementary file 4: Figure 4A).

Merodon opacus Vuji¢, Likov & Radenkovic¢ sp. nov.
http://zoobank.org/256F1010-GAAF-406A-830B-4F4A1D95126A
Figs 12A—F, 13A, B, 14H-J, 20C, 21A—D, 31A, B, 32A—C

Diagnosis. Medium sized (7.2-10.6 mm), short pilose dark species with olive-brown
reflection; antennae dark; legs mostly black; basoflagellomere elongated (1.8—2 times
as long as wide) obviously concave dorsally; arista short 1.5 times as long as basoflagel-

122 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

lomere (Fig. 12A—F); terga dark (Fig. 21A, B); metafemur incrassate covered with very
short pilosity (Fig. 13A, B); male genitalia: posterior surstyle lobe with small lateral
hump (Fig. 14H, I: bp); apical part of anterior surstyle lobe rhomboid (Fig. 14H: al);
lingula medium sized (Fig. 14J: 1). Similar to Merodon serrulatus from which it differs
in dark tergum 2 (in M. serrulatus with small pale lateral maculae). Related to M. hir-
sutus from which can be distinguished by frons covered with dense microtrichia (Fig.
20C) (mostly shiny in MZ. Airsutus), shorter dorsolateral and ventral pile on metafemur
(Fig. 13A, B), and shorter and more adpressed pile on terga in females (Fig. 21D).
Morphologically related to M. defectus sp. nov. from which can be distinguished by
dark tergum 2 (in VM. defectus sp. nov. tergum 2 with yellow-orange lateral maculae).
Additionally, differs from M. defectus sp. nov. by posterior surstyle lobe with developed
lateral hump (Fig. 14I: bp), reduced in M. defectus sp. nov. (Fig. 14B: bp).

Description. Male. Head. Antennae black to dark brown; basoflagellomere elon-
gated 1.8—2 times as long as wide, and 2.3 times as long as pedicel, concave dorsally
with acute apex; large fossette dorsomedial and dorsolateral (Fig. 12A—C); arista dark
and thickened at basal one third, covered with dense microtrichia, 1.5 times as long as
basoflagellomere; face and frons black with gray microtrichia, face covered with dense
whitish gray, and frons with yellowish gray pile; oral margin microtrichose with shiny
lateral areas; lunule shiny black, bare; vertex covered with microtrichia (Fig. 20C);
vertex isosceles, with long, pale whitish yellow pile, in some cases mixed with few black
pile on the ocellar triangle; ocellar triangle equilateral; eyes covered with dense pile;
occiput with gray-yellow pile, covered with a dense, gray microtrichia; eye contiguity
ca. 10-12 facets long; vertical triangle: eye contiguity: frons = 1.2: 1:2.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile; scutum at wing basis with short black pile; scutum with two or more mi-
crotrichose vittae, anteriorly connected and posteriorly reaching the scutellum; scutum
dull; posterodorsal part of anterior anepisternum, posterior anepisternum (except an-
teroventral angle), anterior anepimeron, dorsomedial anepimeron, and posterodorsal
and anteroventral parts of katepisternum with long, pale yellow pile and grayish mi-
crotrichia; wings entirely covered with microtrichia; wing veins brown; calypteres yel-
lowish; halteres yellowish, in some specimens with darker capitulum; legs mostly black,
except brown tarsi ventrally in some specimens; pile on legs pale yellow; metafemur
moderately incrassate, 3.5 times longer than wide; pile on postero- and anteroventral
surface very short with few sparse pile, and ca. as one fourth of width of metafemur,
approximately the same length as pile on dorsal surface (Fig. 13A).

Abdomen. Tapering, 1.2 times longer than mesonotum; terga dark brown to black;
terga 2—4 each with a pair of white microtrichose, wide, oblique fasciae (on tergum
2 more triangular); pile on terga all yellow (Fig. 21A, C); sterna dark brown, covered
with long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid in shape, 1.5 times
longer than wide, covered with dense, short pile (Fig. 14H: al); posterior surstyle lobe
oval with small basolateral protrusion (lateral hump) (Fig. 14H, I: bp); hypandrium
sickle-shaped, without lateral projections; lingula medium sized (Fig. 14J: 1).

Revision of the Merodon serrulatus group 123

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: antennae with rounded tip, basoflagellomere ca. two times
longer than wide, fossette dorsal (Fig. 12D—F); frons with broad microtrichose vittae
along eye margins; frons covered with pilosity of variable color, from mostly gray-yel-
low to predominantly black; ocellar triangle covered with black pile; terga pale pilose,
in some specimens terga 2—4 medially with short adpressed black pile; microtrichose
fasciae on terga 3 and 4 narrower (Fig. 21B, D).

Morphological description of the puparium (Fig. 31A). Length: 7.5 mm, width:
4 mm; light brown in color; sub-cylindrical; rough integument with larval segmenta-
tion persisting as transverse folds and wrinkles; integument covered with small domes
and spicules; pronounced segmental sensilla, bearing seta. The dorsal surface of the
prothorax with a pair of anterior spiracles, which are more than two times longer than
broad at the base, sclerotized, cylindrical in shape, brown in color, apex with two linear
spiracular openings (Fig. 32B). On the anal segment, two different pairs of lappets
present: the ventro-lateral pair represented by fleshy papilla with one sensilla bearing
a seta and the dorso-lateral pair with a very poorly developed basal papilla, apically di-
vided bearing one sensilla with a long seta on top of each division. Cephalopharyngeal
skeleton (Fig. 31B). Robust mandibles with dark highly sclerotized hooks, without
accessory teeth, fused to the external mandibular lobes; the dorsal cornu narrowed,
representing almost the whole length of the ventral cornu. Clypeal sclerite sclerotized,
tentorium and intermediate sclerites highly sclerotized and apparently fused; ventral
cornu elongated and narrow in profile view, wider and more heavily sclerotized at the
posterior end, forming the grinding mill of pestle and mortar construction at the pos-
terior end of the cibarium; cibarium at the base, with a clearly sclerotized end. Posteri-
or respiratory process (Fig. 32A). Brownish, wider than long, very short (barely visible
from dorsal view), button shape, base only slightly wider than apex; dorsal and lateral
surface covered by a barely visible ornamentation resembling a network. The outline of
the spiracular plate sub-elliptical and barely irregular. Spiracular plate with four pairs
of sinuous spiracular openings (clearly separated from each other) around two central
scars, first pair clearly shorter than the others; three or four very small circular nodules
on each side of the surface of spiracular plate in the area of spiracular openings; four
pairs of branched inter-spiracular setae emerging on the outward edges of the spiracu-
lar plate. Pupal spiracles (Fig. 32C). Sclerotized, brownish in color, stout, cylindrical
in shape, almost as long as broad, slightly tapered, with not heavily rounded promi-
nence at the end (length 0.3 mm) separated by a distance of ca. five times their length.
Upper two-thirds of the lateral sides (except for the granularly surfaced apex) covered
with irregularly-spaced, oval-shaped domed tubercles, leaving a more or less triangular
central area free of tubercles on both ventral and dorsal surfaces; 3—7 radially-arranged
spiracular openings on each tubercle. The whole spiracle surface (from the base to the
apex) reticulated with a polygonal pattern, more irregular on ventral side, with poly-
gons being noticeably smaller in the apical part. Material examined. Greece, Lesvos
Island, Agiassos, C. Pérez-Bafén leg.: 1 larva (L3 instar) buried in the ground of a
chestnut forest, 2 Mar 2006; reared, pupa 4 Mar 2006, adult emerged 21 Mar 2006.

124 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Figure 31. Light micrographs of Merodon opacus sp. nov. puparium. A puparium in dorsal view B ceph-
alopharyngeal skeleton in lateral view. Abbreviations: C—cibarium, Cs—clypeal sclerite, Dc—dorsal cornu,
Is—intermediate sclerite, M—mandibles, Mr—mortar, P—pestle, T—tentorium, Vc—ventral cornu. Scale bars:

3 mm (A); 500 um (B).

Etymology. Latin adjective opacus (opaque, not transparent), pertains to the dark
tergum 2, without reddish yellow lateral maculae.

Distribution. Merodon opacus sp. nov. has been recorded on the Greek island of
Lesvos and in western Turkey (Fig. 7).

Revision of the Merodon serrulatus group 125

polar view showing the spiracular plate B anterior spiracle C pupal spiracle. Scale bars: 200 um (A); 100
um (B, C).

Ecology. Preferred environment: forest/open ground; thermophilous and ev-
ergreen Quercus forest; Castanea forest, dry Pinus forest; unimproved grassland and
tracksides (Fig. 35A). Flowers visited: Ornithogalum spp. and Potentilla spp. Flight
period: March-September.

Type material. Holotype. GreEce * 4; Lesvos; Polichnitos; 39°05'02"N, 26°09'13"E;
30 Apr. 2008; A. Vuji¢ leg.; FSUNS 03758. Original label: “HOLOTYPE of Merodon |
opacus Vujic, Likov et / Radenkovié sp.n. 2019” [red label], “Greece, Lesvos, / Polichnitos
30.IV 2008. / Leg. A. Vujic”, “03758” (See Supplementary file 4: Figure 4B). Paratypes.
GREECE, Lesvos * 1 @; Ag. Ermogenis; 39°01'07"N, 26°32'44"E; 2 May 2008; A. Vuji¢
leg.; FSUNS 03760 ® 1 3; Neochori II; 39°01'10"N, 26°20'02"E; 2 May 2016; A. Vujic,
J. Aéanski leg.; FSUNS 11396 ¢ 1 9; Agiassos; 39°03'00"N, 26°22'60"E; 6 Jun. 2004;
M. Kapsali leg.; MAegean * 3 9 9; Agiassos; 39°03'00"N, 26°22'60"E; 23 May 2004; A.
Kyriakopoulos leg.; MAegean * 1 C; same data as for preceding; 24 May 2001; MAegean
¢ 1 9; same data as for preceding; MAegean * 1 4; Agiassos; 39°03'00"N, 26°22'60"E;
Mar. 2006; C. Pérez-Bafién leg.; CEUA * 4 dd; same data as for preceding; 11 Jun.
2005; CEUA * 4 34; same data as for preceding; 10 Jun. 2005; CEUA * 8 99; same
data as for preceding; CEUA * 4 9 9; same data as for preceding; 6 Jun. 2005; CEUA ¢ 4
S44; same data as for preceding; CEUA ¢ 1 3; Agiassos; 39°03'09"N, 26°22'57"E; 860 m

126 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

a.s.l.; 23 May 2004; M. Kapsali leg.; MAegean ¢ 2 S34; 3.5 km S Agiassos; 39°03'09"N,
26°22'57"E; 860 ma.s.l.; 23 May 2004; M. Kapsali leg.; FSUNS 02487, 02503 © 1 33 3.5
km S Agiassos; 39°03'09"N, 26°22'57"E; 860 m a.s.l.; 23 May 2004; A. Kyriakopoulos
leg.; FSUNS 03762 ¢ 1 9; 3.5 km S Agiassos; 39°03'09"N, 26°22'57"E; 860 m a.s.l.; 23
May 2004; M. Kapsali leg.; FSUNS 03764 * 3 29; Agiassos; 39°03'17"N, 26°23'50"E;
760 m a.s.l.; 10 Jun. 2004; M. Kapsali leg; MAegean * 1 C; same data as for preceding;
24 May 2004; MAegean * 2 4; same data as for preceding; A. Kyriakopoulos leg.; MAe-
gean * 1 2; same data as for preceding; MAegean * 2 29; same data as for preceding;
10 Jun. 2004; MAegean * 1 9; 3.8 km SSE Agiassos; 39°03'17"N, 26°23'50"E; 760 m
a.s.l., 10 Jun. 2004; A. Kyriakopoulos leg.; FSUNS 03763 ¢ 1 9; Agiassos; 39°03'45"N,
26°23'30"E; 700 maz.s.l.; 20 May 2004; A. Kyriakopoulos leg.; MAegean * 1 9; same data
as for preceding; 6 Jun. 2004; M. Kapsali leg.; MAegean ¢ 1 4; Agiassos; 39°03'92"N,
26°22'87"E; 27 May 2009; M. Taylor leg.; MZH http://id.luomus.fi/GJ.1133 * 1 4; Agi-
assos; 39°04'09"N, 26°23'17"E; 600 m a.s.l.; 15 May 2004; T. Petanidou leg.; MAegean
546; Agiassos; 39°04'17"N, 26°22'22"E; Sep. 2009; A. Vuji¢ leg.; FSUNS C64, C65,
Z9 to Z11° 10 29; Agiassos; 39°04'17"N, 26°22'22"E; Sep. 2009; A. Vuji¢ leg.; FSUNS
Z12 to Z19, Z28, Z29 * 2 JQ; Agiassos; 39°04'17"N, 26°22'22"E; 8 Jun. 2009; G.
Stahls leg.; MZH GJ.1139, GJ.1141* 4 9 9; same data as for preceding; MZH GJ.1135
to GJ.1138 * 4 44; same data as for preceding; 25 May 2009; MZH GJ.1140, GJ.1142,
GJ.1143, GJ.1145 «1 6; Agiassos; 39°04'25"N, 26°22'35"E; 8 May 2007; G. Stahls
leg.; MZH GJ.1144 * 1 9; same data as for preceding; MZH GJ.1126 ¢ 5 44; same
data as for preceding; 30 May 2009; MZH GJ.1119 to GJ.1123 © 2 29; same data as
for preceding; MZH GJ.1124, GJ.1125¢ 1 CS: same data as for preceding; 27 May 2009;
MZH #1 @; 39°10'17"N, 26°18'14"E; FSUNS 02504 ¢ 1 239910 IZ N, 26 b8 14":
FSUNS 02505 * 3 99; 39°10'17"N, 26°18'14"E; 4 Jun. 2012; A. Vujié, L. Likov leg.;
FSUNS G1747 to G1749 © 1 4; Vatousa; 39°13'51"N, 26°01'23"E; 200 m a.s.l.; 28
May 2001; FSUNS ¢ 1 9; 2.5 km S Gavathas; 39°14'54"N, 25°58'60"E; 28 Apr. 2010;
M. Hull leg.; WML 05042 ¢ 6 O4; near Sikaminea; 39°21'14"N, 26°17'56"E; 11 May
2009; G. Stihls leg; MZH GJ.1127 to GJ.1129, GJ.1132, GJ.1134, GJ.1147 » 2 29;
same data as for preceding; 2 Jun. 2009; MZH GJ.1130, GJ.1131 * 2 99; 5.7 km NW
Mantamados; 39°21'19"N, 26°17'52"E; 600 m a.s.l.; 2-10 May 2001; FSUNS 02488,
02507 * 3 63; Mantamados; 39°21'19"N, 26°17'52"E; 600 m a.s.l.; 10 May 2001;
FSUNS ¢ 2 29; same data as for preceding; FSUNS * 3 43; same data as for preceding;
17 May 2001; FSUNS © 1 9; same data as for preceding; FSUNS ¢ 1 C; same data as for
preceding; 23 May 2001; FSUNS ¢ 1 ; Sikaminia; 39°21'42"N, 26°17'47"E; 10 May
2001; C. Pérez-Bafin, S. Rojo leg.; CEUA * 3 4; same data as for preceding; 14 May
2001; CEUA © 15 34; same data as for preceding; 17 May 2001; CEUA ¢ 1 9; same
data as for preceding; CEUA * 2 34; Sikaminia; 39°21'44"N, 26°17'49"E; 3 May 2008;
A. Vuji¢ leg.; FSUNS 03757, 03759 © 1 G; Sikaminia; 39°21'44"N, 26°17'49"E; 2 May
2001; FSUNS 02506 * 1 9; Near Lepetimnos; 39°21'47"N, 26°16'32"E; 1 May 2016;
A. Vuji¢, J. Acanski leg.; FSUNS 11243.

Turkey ¢ 3 29; 12 km SW of Muégla; 37°07'40"N, 28°16'28"E; 660 m a.s.l.; 23
May 2011; M. Bartak, Kubik leg.; M. B. coll. 17921 to 17923 © 26 Jb; Mugla, Uni-

Revision of the Merodon serrulatus group 127

A B C

Figure 33. Merodon trianguloculus sp. nov., antenna, lateral view. A outer side, male B inner side, male

outer side, female. Scale bar: 1 mm.

versity Campus; 37°09'42"N, 28°22'13"E; 700 m a.s.l.; 17-22 May 2011; M. Bar-
tak, Kubik lees: MB. coll: 179279917928 17930," 179315 1793717938, 91 7940.10
17943, 17946, 17951, 17955 to 17960, 17965 to 17972 © 20 99; Mugla, University
Campus; 37°09'42"N, 28°22'13"E; 700 ma.s.l.; 17-22 May 2011; M. Bartak, Kubik
leg.; M. B. coll. 17929, 17932 to 17936, 17939, 17944, 17945, 17947 to 17950,
17952 to 17954, 17961 to 17964 © 1 2; Mugla, University campus; 37°09'42"N,
28°22'21"E; 700 m a.s.l.; Apr—May 2014; O. Dursun leg.; M. B. coll. 10463 © 3
29; Mugla, 13 km NE pine wood; 37°14'50"N, 28°30'00"E; 1200 m a.s.1.; 23-27
Jun. 2015; M. Bartak, Kubik leg.; M. B. coll. 17924 to 17926 ¢ 4 99; Bozdag moun-
tain, Near Bozdag; 38°22'28"N, 28°04'38"E; 1140 ma.s.l.; 7 Jun. 2014; A. Vujié, J.
Aéanski leg.; FSUNS 06945, 06946, 06948, 06951 ¢ 2 Jb; Bozdag mountain, Near
Bozdag; 38°22'28"N, 28°04'38"E; 1140 ma.s.l; 7 Jun. 2014; A. Vuji¢, J. Acanski
leg.; FSUNS 06947, 06949.

Other material. Greece, Lesvos * 1 9; Agiassos; 39°03'00"N, 26°22'60"E; 28
May 2010; Horsfield leg.; NMS ¢ 1 9; Agiassos; 39°03'00"N, 26°22'60"E; 3 Jun.
2010; Wilkinson leg.; NMS [published in Ricarte et al. (2012), as Merodon serrulatus]
¢ 1 9; same data as for preceding; 8 Jun. 2010; NMS ¢ 1 9; Agiassos; 39°03'00"N,
26°22'60"E; 1-8 Jun. 2010; Hancock leg.; GLAHM ¢ 2 4; same data as for preced-
ing; 28-29 May-Jun. 2010; GLAHM ° 2 &; Agiassos; 39°03'36"N, 26°23'30"E; 1
Jun. 2010; Horsfield leg.; NMS [published in Ricarte et al. (2012), as Merodon serrula-
tus] ° 1 9; Agiassos; 39°03'36"N, 26°23'30"E; 28 May 2010; Horsfield leg.; NMS ¢ 1
3S; 3.5 km § Agiassos; 39°04'15"N 26°22'17"E; 860 m a.s.L.; 8 Jun. 2004; M. Kapsali
leg.; MZH ¢ 1 3; Agiassos; 39°04'17"N, 26°22'22"E; 1910 ma.s.l.; 26 May 2003; C.
J. Palmer leg.; WML 298/03 © 1 9; same data as for preceding; WML 310/03 ¢ 1 3;
SW from Agiassos; 39°04'35"N, 26°22'03"E; 24 May 1988; NBCN ° 1 Q; Potamia
river; 39°13'36"N, 26°06'48"E; 1-8 Jun. 2010; Hancock leg.; GLAHM » 1 S;2km S
Gavathas; 39°16'11"N, 25°58'33"E; 5 May 2005; M. Hull leg; WML ° 1 3; 5.7 km
NW Mantamados; 39°21'19"N, 26°17'52"E; 31 May 2004; E. Lamborn leg.; MZH.

128 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Merodon sacki (Paramonov, 1936)
Figs 8C, 9D-F, 10C, D, 11D-F

Diagnosis. Large (9.5—11.6 mm) dark brown species with lack of microtrichose fasciae
on terga 2—4 in males (Fig. 10C) and curved and very incrassate metafemur with long
pile on ventral margin; the longest pile as long as half of width of metafemur (Fig. 8C).
Similar to Merodon bequaerti but differs by strongly curved metafemur and generally
longer body pile, clearly visible on tergum 4 (Fig. 10D).

Redescription (based on holotype and additional material from the type area,
Spain). Male. Head. Antennae black to dark brown; basoflagellomere ca. two times
as long as wide, and ca. two times as long as pedicel, concave dorsally; large fossette
dorsolateral; arista dark and thickened at basal one third, covered with dense micro-
trichia, 1.6 times as long as basoflagellomere (Fig. 11D-—F); face and frons black with
gray microtrichia, face covered with dense whitish gray, and frons with yellowish gray
pile; oral margin microtrichose with shiny lateral areas; lunule shiny black, bare; vertex
covered with golden microtrichia around ocellar triangle; vertex isosceles, with long,
pale whitish yellow pile mixed with black pile on the ocellar triangle; ocellar triangle
equilateral; eyes covered with dense pile; occiput with gray-yellow pile, covered with a
dense, gray microtrichia; eye contiguity 10-14 facets long.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile; scutum at wing basis with short black pile; scutum with two or more mi-
crotrichose vittae, anteriorly connected and posteriorly reaching the scutellum; scutum
dull; posterodorsal part of anterior anepisternum, posterior anepisternum (except an-
teroventral angle), anterior anepimeron, dorsomedial anepimeron, and posterodorsal
and anteroventral parts of katepisternum with long, pale yellow pile and grayish micro-
trichia; wings entirely covered with microtrichia; wing veins brown; calypteres and hal-
teres pale yellowish; legs mostly black, except brown tarsi ventrally in some specimens;
pile on legs pale yellow, except black pile at apical one fourth of metafemur; metafemur
curved and incrassate, approximately three to four times longer than wide; pile on
postero- and anteroventral surface long, and ca. half of width of metafemur (Fig. 8C).

Abdomen. Broad, tapering, 1.2 times longer than mesonotum; terga dark brown
to black, usually without microtrichose fasciae; tergum 2 with orange lateral maculae;
pile on terga all yellow (Fig. 10C, D); sterna dark brown, covered with long whitish
yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid in shape, 1.5 times
longer than wide, covered with dense, short pile (Fig. 9D: al); posterior surstyle lobe
oval with basolateral protrusion (lateral hump) (Fig. 9D, E: bp); hypandrium sickle-
shaped, without lateral projections; lingula large (Fig. 9F: 1).

Female. Unknown.

Distribution. Merodon sacki is known only from Spain (Fig. 7).

Ecology. Preferred environment: forest/open ground; open areas in evergreen oak
forest (Quercus ilex and Q. suber) and Mediterranean scrub. Flowers visited: no data.

Flight period: April-July.

Revision of the Merodon serrulatus group 129

Type material. Holotype (original designation): male, “Holotypus Lampetia /
sacki Paramononv, 1936 / G.V. POPOV des. 2007” [red label], “Lampetia | sacki n. sp.
! 3 Typus | Paramonov d.” [pink label handwritten], “Merodon | mir tinbekannt” [yel-
low label handwritten], “74 VIT 81” “Chiclana’ |handwritten on the back side] (SIZK)
(See Supplementary file 6: Figure 6B) (studied).

Note (Popov pers. comm.). The species was described by examining a single male,
with the type clearly indicated on the label by Paramonov (discovered and deposited
in SIZK). The type specimen is considered lost (Liepa 1969). Hurkmans (1993: 178,
179) incorrectly considered M. sacki as a junior synonym of M. clavipes (Fabricius,
1781). Hurkmans (1993) also provided an incorrect year for the description of Lam-
petia sacki (1937 instead of the correct 1936), and also incorrectly designated the lec-
totype and paralectotype [Articles 73 and 74 of the ICZN (1999)] for two M. clavipes
females with the same label “Chiklana’, which are not syntypes [a violation of Articles
74.1 and 74.2 of ICZN (1999)]. The holotype was established by the original designa-
tion according to Article 73.1.1 of the ICZN (1999), as well as by a monotype accord-
ing to Article 73.1.2 (ibid.).

Other material. Spain * 3 JG; La Corte; 37°57'41"N, 6°49'09"W; 28 Apr. 2015;
A. Vuji¢, D. Obreht leg.; FSUNS 09340, 09343, 09345.

Merodon serrulatus (Wiedemann in Meigen, 1822)
Figs L[A—G, 2A—D, 3A-J, 4A-F, 5A—D, 6A-G, 14C

Merodon alexeji Paramonov, 1925: 155 — syn. published in Vuji¢ et al. 2011: 84.

Merodon lusitanicus Hurkmans, 1993: 181 — syn. published in Marcos-Garcia et al.
2007: 566.

Merodon tener Sack, 1913: 443 syn. nov.

Diagnosis. Medium sized (7.1—-10.9 mm), short pilose dark species with olive-brown
reflection; antennae dark brown; legs mostly black; body pile predominantly pale yel-
low, except black pile on vertex and scutum, terga 2—4 in some specimens and apical
one third of femora in some specimens and populations; basoflagellomere elongated
(1.7—2.2 times as long as wide) obviously concave dorsally, arista short (Fig. 3); tergum
2 usually with small pale orange-yellow lateral maculae (Fig. 2); metafemur incrassate,
ca. three times longer than wide, with short pilosity, except few long pile on postero-
and anteroventral surface of metafemur (Fig. 4); male genitalia: apical part of anterior
surstyle lobe triangular (Fig. 1A: al, C, D); posterior surstyle lobe with lateral hump
(Fig. 1A, B: Ip); lingula large (Fig. 1E: 1).

Redescription (based on the types and specimens from the type area of nominal
taxon, Iberian Peninsula; variability includes populations from all of the range). Male.
Head. Antennae black to dark brown; basoflagellomere (Fig. 3A—C, G—J) elongated,
1.7—2.2 times as long as wide, and 2.5—3 times as long as pedicel, concave dorsally,
tapering to the apex; dorsolateral and dorsomedial (if present) fossette large with vari-

130 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

able shape (see variability) (as on Fig. 3); arista dark and thickened at basal one third,
covered with dense microtrichia; arista short, 1.2—1.5 times as long as basoflagellomere
(Fig. 3); face and frons black with gray microtrichia, face covered with dense whitish,
and frons with yellowish gray pile; oral margin shiny, with small lateral microtrichose
area; lunule shiny black, bare; eye contiguity 8-10 facets long; vertex isosceles, shiny
black, except in front of anterior ocellus, covered with microtrichia; vertex with long,
pale whitish yellow pile, in some cases mixed with few black pile on the ocellar triangle;
ocellar triangle from equilateral to isosceles (see variability); occiput with gray-yellow
pile, ventrally covered with a dense, gray microtrichia; eyes covered with dense whitish
pile (Fig. 5A, B); ratio of length of vertical triangle: eye contiguity: frons = 3: 1 : 3.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
usually yellow pile; scutum at wing basis in some specimens and populations with
patch of black pile, or with fascia of black pile between wing basis; scutum usually with
two or four microtrichose vittae (see variability), anteriorly connected and posteriorly
reaching the scutellum (Fig. 2A); anterior half of scutum from dull until shiny black
(see variability); posterodorsal part of anterior anepisternum, posterior anepisternum
(except anteroventral angle), anterior anepimeron, dorsomedial anepimeron, and pos-
terodorsal and anteroventral parts of katepisternum with long, dense pale yellow pile
and grayish microtrichia; wings entirely covered with microtrichia; wing veins brown;
calypteres pale yellow; halteres yellow, in some cases with dark capitulum; legs (Fig. 4)
without spinae or other protuberances; legs mostly black, except brown tarsi ventrally
in some specimens; pile on legs pale yellow, except black pile at apical one third of
metafemur in some populations (see variability); metafemur moderately incrassate, ca.
three times longer than wide; long pile on postero- and anteroventral surface sparse,
and ca. one third to one fourth (see variability) of width of metafemur, approximately
the same length as pile on dorsal surface (Fig. 4).

Abdomen. Tapering posteriorly, ca. 1.2 times longer than mesonotum; terga dark
brown to black, except for a pair of pale yellow-orange, triangular, lateral maculae on
tergum 2 (in some specimens less visible: see variability); terga 3 and 4 each with a pairs
of white microtrichose, oblique fasciae (on tergum 2 triangular); color of pile on terga
variable, from all yellow to specimens with many black pile on terga 2—4 (see variabil-
ity) (Fig. 2C, D); sterna dark brown, covered with long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe triangular shape, 1.1—1.4 times
longer than wide, covered with dense, short pile (Fig. 1A: al, C, D); posterior surstyle
lobe oval with basolateral protrusion (lateral hump) (Fig. 1A, B: bp, 14C: bp); cercus
rectangular (Fig. 1A: c); hypandrium sickle-shaped, without lateral projections; lingula
large (Fig. 1E: 1).

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: antennae with rounded tip, basoflagellomere ca. two times
longer than wide, fossette dorsal (Fig. 3D—F); frons with microtrichose vittae along eye
margins variable in shape and size (see variability); frons covered with variable color of
pilosity, from mostly gray-yellow until predominately black (see variability) (Fig. 5C,
D); ocellar triangle covered with black pile; lateral side of terga, anterior two third of

Revision of the Merodon serrulatus group 13

tergum 2 and all tergum 5 with yellow pile; central part of terga 2-4 with short ad-
pressed black pile; microtrichose fasciae on terga 3 and 4 conspicuous (Fig. 2D).

Variability. There is some intra- and interpopulation variability in the morpho-
logical characters of Merodon serrulatus, which are summarized in Table 1.

Distribution. As shown in Fig. 7, this Merodon taxon is characterized by the great-
est range, extending from Iberian Peninsula in the south-west, through Greece and east-
ern Turkey to the south, and eastward to Siberia and Mongolia (Doczkal pers. comm.).

Ecology. Preferred environment: forest/open ground; thermophilous Quercus for-
est; Castanea forest (Stahls et al. 2009), evergreen oak forest (Quercus ilex L. and Q.
suber L.), dry Pinus forest; lentisc scrub; dry, well-vegetated, calcareous and non-calcar-
eous unimproved grassland and tracksides; hedgehog heath (Speight 2018); Pinus, Pi-
cea, and Larix forests (Siberia) (Fig. 35B). Flowers visited: Umbelliferae; Cirsium spp.,
Helianthemum spp., Potentilla spp., Rosa spp., Thapsia spp., and Thymus spp. (Speight
2018). Flight period: April-August.

Type material. Holotype of Merodon serrulatus [original designation in Meigen
(1822: 360)]: Wiedemann in Meigen (1822) as Lampetia serrulata: “Portugal / Hoff-
mannsegg S.” 1 2, (ZHMB) (studied).

Merodon alexeji: Described by Paramonov (1925) based on two specimens (male
and female). Lectotype [designated by Marcos-Garcia et al. (2007)]: male, “Merodon /
alexeji n. sp. | Typus / Paramonov d.”, “Kohanovka / Baltsk. u. / Odes. g. [in Cyrillic]
1.VI.24. Ucraina’” (PC) (SIZK) (studied).

Merodon lusitanicus: Holotype [original designation by Hurkmans (1993: 181)]:
female, “Portugal, Algarve, Quarteira 27.iv.1985, J.A.W. Lucas” (NBCN) (studied).
Paratypes. Portucat * 1 Q; Algarve, Quarteira; 37°03'29"N, 8°04'47"W; 27 Apr.
1985; NBCN « 1 9; Algarve, Vilamoura; 37°04'35"N, 8°07'46"W; 27 Apr. 1985;
NBCN.

Merodon tener: Described by Sack (1913: 443) based on three male and three fe-
male syntypes. Lectotype [designated by Hurkmans (1993)]: female “Sarepta [ = Kras-
noarmeysk near Volgograd, after Peck 1988] / M. tener Sack det. Sack / coll. Lichtwardt
/ coll. D. E. I. Eberswalde” (ZHMB) (studied). Original label: “LECTOTYPE of /
M. tener Sack / des. 1988 Hurkmans” [red label handwritten], “Sarepta” [yellow label
handwritten], “M. tener Sack / det. Sack” [label partly handwritten], “Coll. DEI /
Eberswalde”, “Coll. Lichtwardt”. (See Supplementary file 6: Figure GA). Lectotype is
conspecific with type of M. serrulatus, sharing the same morphological characters.

Other material. Croatia ¢ 1 9; Velebit, BruSane; 44°29'55"N, 15°16'43"E; 600
ma.s.l.; 13 Jun. 1969; NBCN 02489.

France * 1 9; Languedoc-Roussillon, Corbieres, Carcassonne; 43°13'00"N,
2°21'00"E; 18 Jun. 1974; NBCN ° 1 3; Provence Alpes Cote d’Azur, Saint-Maximin-
la-Sainte-Baume; 43°25'23"N, 5°50'11"E; 17-20 Jun. 1951; M. Bequaert leg.; NBCN
02491 © 1 9; same data as for preceding; NBCN 02492 « 1 9; Source du Lez, Saint
Clement; 43°43'05"N, 3°50'39"E; 24 May 1989; Maldes leg.; MNHN 22629 1 4;
Provence Alpes Cote d'Azur, Montagne du Luberon, W from Bonnieux; 43°48'00"N,
5°22'00"E; 3 Jun. 1993; NBCN ¢ 1 9; Feuilla, Route de Treilles, en face du village Panais;

132: Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

43°53'53"N, 2°00'16"E; 7 Jun. 1988; J. Hamon leg.; MNHN 17973 * 3 3; Departe-
ment du Gard, Mas Mejean; 44°05'24"N, 3°35'26"E; 29 May 1952; NBCN e 2 JSS; P
N. Mercantour, Le Bor, on, Umgebung, mesophiles pot. Argent; 44°06'47"N, 7°16'42"E;
1380 ma.s.l.; 21 Jun. 2011; A. Ssymank leg.; A. S. coll. G1057 © 1 CS; Causse de Sauvet-
erre; 44°22'03"N, 3°13'49"E; 20 Jul. 1971; MNHN 22628 ¢ 1 9; Larche (Basses Alpes);
44°26'59"N, 6°50'60"E; 22 Jul. 1925; R. Benoist leg; MNHN PM0383 « 1 ; same
data as for preceding; MNHN PM0429 « 1 CS; same data as for preceding; 3 Jul. 1925;
MNHN 22630 © 1 @; Provence Alpes Cote d’Azur, Larche; 44°26'59"N, 6°50'60"E;
22 Jul. 1923; NBCN ¢ 1 4; Drome, La Chapelle en Vercors; 44°58'12"N, 5°23'39"E;
28 Jun. 1970; Roman Emile leg; MNHN PM0377 ¢ 2 44; Isere, Villars de Lans Pic st
Michel hill top; 45°05'24"N, 5°37'12"E; 1970 m a.s.l.; 20 Jul. 2010; J. van Steenis leg.;
J. v. S. coll. © 1 Q; LArselle; 45°23'10"N, 7°04'19"E; 14 Jun. 1909; MNHN PM0357.
GREECE ¢ 2 9 9; Mountain Taygetos; 22 km SW Sparta; 36°58'60"N, 22°24'09"E;
6 May 1990; NBCN ¢ 1 4; Mountain Taygetos; 37°05'20"N, 22°18'55"E; 950-1800
m a.s.l.; 15-19 May 1990; ZMUC 00513256 * 7 3; Laconia, Karyes, 25 km N
from Sparta; 37°18'15"N, 22°25'16"E; 930 ma.s.l.; 23 May 2014; A. Vuji¢, J. A¢anski
leg.; FSUNS 06535, 06542, 06547, 06549, 06556, 06563, 06560 * 6 QQ; Laco-
nia, Karyes, 25 km N from Sparta; 37°18'15"N, 22°25'16"E; 930 m a.s.l.; 23 May
2014; A. Vujic, J. Acanski leg.; FSUNS 06543, 06544, 06546, 06553, 06555, 06565
° 3 Oo; Chelmos, Kalavryta ski center; 38°00'25"N, 22°11'40"E; 6 Jun. 2017; A.
Vujic, Z. Nedeljkovi¢é, L. Likov, M. Milici¢é, T. Tot, leg.; FSUNS 15980 to 15982
¢ 2 QQ; Chelmos, Kalavryta ski center; 38°00'25"N, 22°11'40"E; 6 Jun. 2017; A.
Vujic, Z. Nedeljkovic, L. Likov, M. Milici¢, T. Tot leg.; FSUNS 15983, 15984 ¢ 1
@; Achaia, Mountain Chelmos above Kalavryta; 38°00'31"N, 22°07'08"E; 1700 m
a.s.l.; 17-19 Jun. 1982; B. Skule, S. Langemark leg.; ZMUC 00513264 * 3 99;
Achaia, Mountain Chelmos above Kalavryta; 38°00'31"N, 22°07'08"E; 1700 m
a.s.l.; 17-19 Jun. 1982; B. Skule, S. Langemark leg.; ZMUC 00513265, 00513273,
00513301 © 2 64; Corfu; 39°40'00"N, 19°45'00"E; NHMW 02485, 02486 © 2 4;
Corfu; 39°40'00"N, 19°45'00"E; 1400 m as.l.; NHMW 1 9; Peristeri mountain;
39°40'36"N, 21°07'06"E; 2030 m a.s.1.; 24-28 May 1994; V. Michelsen leg.; ZMUC
00513259; © 1 4; 15 km NO Metsovo; 39°47'19"N, 21°11'58"E; 4 Jun. 1994; M.
Ohl leg.; ZHMB ° 4 4; Mountain Pindos, Katara Pass; 39°47'46"N, 21°13'44"E;
20 May 1997; S. Radenkovié leg.; FSUNS ¢ 3 4; same data as for preceding; S.
Simié leg.; FSUNS ¢ 4 2 9; same data as for preceding; A. Vuji¢ leg.; FSUNS * 6 od;
Mountain Pindos, Katara Pass; 39°47'48"N, 21°13'45"E; 1700 ma.s.1.; 20 May 1997;
FSUNS 01779, 01781 to 01785 * 3 2Q; same data as for preceding; FSUNS 01786,
01787, 01780 ¢ 1 2; Mountain Pindos, Katara Pass; 39°54'00"N, 21°11'00"E; 13 Jul.
1979; M. C. D Day, G. R. Else, D. Morgan leg.; NHMUK ¢ 1 &; Mountain Pindos,
“Iznad Panagije” [Panagia]; 39°48'25"N, 21°19'44"E; 850 m a.s.L; 15 May 2011; A.
Vuji¢ leg.; FSUNS H38 ¢ 15 44; Mountain Olympos, Litochoras-Prionia 3, “pro-
planak pored puta”; 40°04'36"N, 22°00'46"E; 17 May 2012; A. Vuji¢ leg.; FSUNS
H82, H83, H86 to H90, H92, H93, H95 to H97, H99, 16, 17 © 1 3; Mountain
Olympos, Litochoras-Prionia 3, “proplanak pored puta’; 40°04'36"N, 22°00'46"E; 17

Revision of the Merodon serrulatus group 133

May 2012; FSUNS H94 © 1 3; Mountain Olympos, Near Litochoro; 40°06'30"N,
22°28'41"E; 21 May 2014; A. Vuji¢, J. Aéanski leg.; FSUNS 06499 ¢ 1 2; Mt Olym-
pos, Litochoro; 40°06'41"N, 22°28'37"E; 650 m a.s.l; 17 May 2016; A. Vuji¢, J.
Aéanski, M. Milici¢, Z. Nedeljkovi¢ leg.; FSUNS 11679 ¢ 8 44; Mountain Olympos,
Litochoras-Prionia 4; 40°06'43"N, 22°28'08"E; 18 May 2011; A. Vuji¢ leg.; FSUNS
I11 to 114, 117, 120 to 122 © 2 99; same data as for preceding; FSUNS 115, I16.

Itaty ¢ 1 9; Sicily, Etna, Rifugio Filiciusa; 37°43'14"N, 15°02'51"E; 1400-1500 m
a.s.l.; 22-28 Jul. 1961; V. S. van der Goot leg.; NBCN ° 1 ; Toscana, Florence, Careggi;
43°48'45"N, 11°15'07"E; 19 May 1986; NBCN °¢ 2 44; Piedmont, Colle di Sestrieres;
44°57'00"N, 6°52'60"E; 1800/2100 m a.s.l.; 23-31 Jul. 1837; Zerny leg.; NHMW.

MONTENEGRO * 1 6; Lovéen, Lovéen 1; 42°22'59"N, 18°53'54"E; 17 May
2018; A. Vujic, A. Sebié, M. Rankovié leg.; FSUNS 19017 © 1 CS; Boka Kotorska,
Morinj; 42°29'25"N, 18°38'56"E; 16-18 May 1998; FSUNS 03600 ° 1 CS; same data
as for preceding; 18-19 May 1998; FSUNS 03601 ¢ 2 99; same data as for preceding;
FSUNS 03602, 03603.

KAZAKHSTAN * 1 9; East Kazakhstan, Markakol’ District, 20 km N settlement
Alekseevka, Souther slop of Matobaj Mountain range; 48°42'22"N, 85°57'00"E; 2318
ma.s.l.; 6 Jul. 1996; V. Zinchenko leg.; SZMN ¢ 1 9; Kazakhstan, 9 km S settlement
Karaoj, Kyzyl-Tass Mountain; 29 Jun. 1996; V. Zinchenko leg.; SZMN.

NORTH MACEDONIA * 1 3; Kozuf, Golema poljana; 41°10'54"N, 22°12'05"E;
15 Jun. 1955; FSUNS 00165 © 1 3; same data as for preceding; 18 Jun. 1956; FSUNS
00166 * 5 4; Kozuf, Golema poljana; 41°10'54"N, 22°12'05"E; 14 Jun. 1975;
FSUNS 00154, 00156 to 00158, 00162 © 1 9; Kozuf, Golema poljana; 41°10'54"N,
22°12'05"E; 15 Jun. 1968; FSUNS 00167 * 2 29; same data as for preceding; 17
Jun. 1956; FSUNS 00168, 00169 * 6 29; Kozuf, Golema poljana; 41°10'54"N,
22°12'05"E; 14 Jun. 1975; FSUNS 00155, 00159 to 00161, 00163, 00164 * 2 99;
Kozuf; 41°25'38"N, 21°30'45"E; 14 Jun. 1975; FSUNS 00152, 00153.

Russia * 1 4; Sarepta, “Russia, RUS, Sarepta, now a suburb of Volgograd city
(Christoph)”; 48°31'40"N, 44°29'01"E; ZHMB 02501 ¢ 4 29; same data as for pre-
ceding; ZHMB 02500, 02502, 02508, 02509 © 32 4; Altai, 10 km S-W of Katanda;
50°06'51"N, 86°07'21"E; 6 Jul. 1983; A. Barkalov leg.; SZMN * 9 29; same data as
for preceding; SZMN » 1 3; SW Altai, Katun valley 10 km W Katanda; 50°09'46"N,
86°06'50"E; 7 Jul. 1983; H. Hippa leg.; MZH * 1 3; same data as for preceding; 22—
27 Jun. 1983; MZH; “exp. Mikkola, Hippa et Jalava” ¢ 1 G; Altai, Kurayskaya Step’;
50°12'00"N, 87°47'60"E; 1662 ma.s.L.; 9 Jul. 2006; A. Barkalov leg.; SZMN ¢ 17 spec-
imens; Altai, Terekta; 50°16'12"N, 85°58'12"E; 1098 m as.l.; Jun. 1973; SZMN « 1
3; Tuva, Erzin river; 50°19'12"N, 95°30'00"E; 1288 m a.s.l.; 27 Jun.—1. Jul. 1989; D.
Logunov leg.; SZMN * 16 2 Q; same data as for preceding; SZMN ¢ 2 43; Tuva, Tere
Khol’ Lake; 50°42'11"N, 97°20'2"E; 27 Jun.—1 Jul. 1989; D. Logunov leg.; SZMN ¢ 1
specimen; Altai, Tuyekta; 50°51'00"N, 85°49'48"E; 944 m a.s.l.; Jun. 1979; SZMN ¢ 2
specimens; Tuva, Chagytay; 50°58'30"N, 94°38'47"E; 1963; SZMN ® 1 specimen; Altai,
Baragash; 51°16'48"N, 85°12'36"E; 927 m a.s.L; Jun. 1973; SZMN ¢ 2 99; Altaj, Er-
lagol; 51°22'21"N, 86°05'22"E; 27 Jun. 1995; FSUNS 00171, 00173 * 3 33; Altaj, Er-

134 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

lagol; 51°22'23"N, 86°05'29"E; 27 Jun. 1995; A. Tepavcevi¢ leg.; FSUNS 00170, 00172,
02476 ° 1 9; same data as for preceding; FSUNS 02477 ° 1 3; Gornyi Altai, Turochaksky
r-n kordon Obogo; 51°35'47"N, 87°05'45"E; 950 ma.s.L; 15 Jun. 2003; D. Kropaceva
leg.; SZMN 22631 ¢ 18 44; Altai mountains, Teletskoye Lake; 51°41'20"N, 87°33'43"E;
23-25 Jun. 2013; A. Vuji¢, S. Radenkovié leg.; FSUNS NJ56, NJ57, NJ59 to NJ62,
NJ64 to NJ71, NJ73 to NJ75, NJ77 * 5 99; same data as for preceding; FSUNS NJ63,
NJ72, NJ76, NJ78, NJ79 * 2 3; Altai, Turochaksky r-n, Teletskoe Lake, 14 km S of
Iogach; 51°42'0"N, 87°17'60"E; 598 m a.s.L; 27 Jun. 2006; V. Zinchenko leg.; SZMN
2 99; same data as for preceding; SZMN ¢ 1 9; Mountain Ural, Orenburg; 51°46'12"N,
54°59'53"E; ZHMB 02519 * 15 GG; Siberia, Altaya, Teletskoe Lake; 51°46'60"N,
87°18'00"E; 24-19 Jun. 2006; A. Barkalov, V. Zinchenko leg.; SZMN ¢ 6 9 9; same data
as for preceding; SZMN ¢ 2 43; same data as for preceding; 27 Jun. 2006; J. T. Smit leg.;
FSUNS 03972, 03973 © 1 9; same data as for preceding; 24 Jun. 2006; FSUNS 03974
¢ 1 &; Siberia, Republic Alatai, Teletskoe lake, Artybash.; 51°47'57"N, 87°14'58"E; 25
Jun. 1990; G. Stahls leg; MZH ¢ 8 44; Altai, Teletskoe Lake, Artibash; 51°47'57"N,
87°14'58"E; 12 Jun. 1990; A. Barkaloy, Cekanov leg.; SZMN ¢ 1 9; same data as for pre-
ceding; SZMN * 17 3G; same data as for preceding; 11-25 Jun. 1990; SZMN * 8 99;
same data as for preceding; SZMN ¢ 13 4; same data as for preceding; 18-20 Jun. 1990;
SZMN ¢ 10 29; same data as for preceding; SZMN ¢ 1 9; same data as for preceding;
23 Jul. 1979; SZMN ° 82 specimens; same data as for preceding; Jun. 1979; SZMN ¢ 1
4; Altaj, Gorno-Altaysk; 51°57'08"N, 85°57'19"E; 21 Jun. 1983; MZH; “exp. Mikkola,
Hippa et Jalava” * 152 specimens; Altai, Gorno-Altaysk; 51°57'08"N, 85°57'19"E; Jun.—
Jul. 1979; A. Barkalov leg.; SZMN ¢ 2 4; same data as for preceding; 22 Jun. 1983;
SZMN ¢ 3 29; same data as for preceding; SZMN ¢ 1 4; Altai Republic; 52°30'00"N,
83°00'00"E; 25 Jun. 1979; NBCN ® 1 9; same data as for preceding; NBCN ¢ 2 J;
Sayan Mountains, Abaza; 52°41'17"N, 90°05'27"E; 30 May—11 Jun. 1981; A. Barkalov,
T. Varlamova leg.; SZMN ° 7 specimens; same data as for preceding; Jun. 1969; SZMN »
2 specimens; Novosibirsk; 55°06'56"N, 82°51'33"E; 1972-1974; SZMN ® 1 specimen;
Tuva, Sosnovka; 56°18'18"N, 51°14'44"E; 1949; SZMN.

SPAIN * 1 @; Sierra Nevada, second valley; 37°06'10"N, 3°27'19"W; 1430 m a.s.L;
17 Jun. 2014; A. Vujic, S. Radenkovi¢, C. Pérez-Bafén leg.; FSUNS 07410 ¢ 3 exer
Sierra Nevada, Ski Centar Sierra Nevada; 37°06'45"N, 3°25'10"W; 2190 m as.L.; 16
Jun. 2014; A. Vuji¢, S. Radenkovié, C. Pérez-Bafién leg.; FSUNS 07275, 07287, 07302
¢ 1 3; prov. Granada Sierra Nevada ri. Valetta; 37°06'55"N, 3°29'32"W; 1 Jun. 1982;
NBCN ° 11 43 Sierra Nevada, First valley; 37°07'40"N, 3°26'44"W; 1630 m a.s.L;
17 Jun. 2014; A. Vuji¢é, S. Radenkovic, C. Pérez-Bafén leg.; FSUNS 07325, 07326,
07328, 07338, 07342, 07344, 07358, 07368, 07371, 07373, 07384 © 10 29; same data
as for preceding; FSUNS 07324, 07340, 07341, 07343, 07355, 07375, 07381, 07385,
07392, 07401 * 4 dC; Sierra Nevada, road to hotel Duque; 37°08'17"N 3°25'46"W;
16 Jun. 2014; A. Vujié, S. Radenkovic, C. Pérez-Bafén leg.; FSUNS 07248, 07251,
07255, 07256 © 3 PQ; same data as for preceding; FSUNS 07262, 07263, 07265
1 4; Sierra Nevada Lugros, Horcajo del Camarate; 37°11'50"N, 3°15'13"W; 1370 m
a.s.l.; 18 Jun. 2014; A. Vuji¢, S. Radenkovi¢, C. Pérez-Bafén leg.; FSUNS 07428 ¢ 1
G; Andalusia, Sierra de Baza, Prados del Roy; 37°22'33"N, 2°51'06"W; 2000/2100 m

Revision of the Merodon serrulatus group 135

Table |. Inter- and intrapopulation variability of Merodon serrulatus.

Character Variability (intra- and/or Intra Inter
interpopulation)
color of antenna from black to brown + -
length of basoflagellomere 1.7—2.2 times as long as wide + + shorter in Balkans populations
(1.7-1.9)
position of antennal fossette| dorsal to lateral or dorsal and medial + + Iberian populations with medial
in male (Fig. 3) fossette
length of arista 1.0-1.5 times as long as basoflagellomere - + longer in Balkans populations
(1.4)
ocellar triangle in male equilateral or isosceles + -
microtrichose vittae on from 2-4, posterior half dull without 1 -
scutum microtrichia
black pile on scutum few, or fascia of black pile between wing + E
basis, or many black pile on scutum
color of pile on metafemur | all yellowish to whitish or with many + + some Iberian populations with
in male black in apical one third only pale pile
length of pile on metafemur one third to one fifth of width of + + in eastern populations (southern
in male metafemur Russia to Siberia) longer, from one
third to one fourth of width of
metafemur
color of knees, apex of from black to brown + -
tibiae and tarsi
lateral maculae on tergum 2 distinct, indistinct, to almost absent + -
pile on terga 3-4 in male from all pale yellow to many black + + Balkans populations with more
black pile
microtrichose vittae on from narrow unconnected to broad and + -
frons in female connected near ocellar triangle
color of pile on frons in almost all black to mostly whitish + -
female
male genitalia the shape of surstyle and size of area 1 + in eastern populations (southern
covered with dense short marginal setulae Russia to Siberia) basolateral
on anterior surstyle lobe (Figs 1, 6) protrusion less distinct (Fig. 6G)

a.s.l.; 9 Jun. 2003; D. Doczkal leg.; D. D. coll. 04805 * 1 9; same data as for preced-
ing; D. D. coll. 04808 ¢ 2 4; Andalusia, Sierra de Baza, Santa Barbara; 37°23'16"N,
2°50'43"W; 1890 ma.s.L; 9 Jun. 2003; D. Doczkal leg.; D. D. coll. 04806, 04807 ¢ 1
9; La Corte; 37°57'41"N, 6°49'09"W; 28 Apr. 2015; A. Vuji¢é, D. Obreht leg.; FSUNS
09333 ¢ 1 &; Alicante, Alcoy-Font Roja; 38°42'00"N, 00°28'00"W; 31 May 1994;
P. M. Isidro leg.; FSUNS 02494 © 1 9; Valensija, Utiel; 39°34'13"N, 1°11'15"W; 9
May 1994; C. Pérez-Bafién leg.; FSUNS 02495 « 1 ; Val de Cabras; 40°09'23"N,
2°01'48"W; 10 Jun. 1980; H. G. M. Tenuissen leg.; NBCN ¢ 1 ; between Leon and
Oviedo, Puerto de Pajares; 43°00'00"N, 5°46'00"W; 12 Jul. 1972; NBCN.

Turkey * 5 Od; “Kop Dags gecidi” [Kop mountain pass], Bayburt; 40°15'00"N,
40°15'00"E; 16 Jul. 1992; NBCN.

Merodon sophron Hurkmans, 1993
Figs 8D, 9G-I, K, 11G-I

Diagnosis. Medium sized (7.8—9.2 mm), dark species with olive-brown reflection;
antennae dark; legs mostly black; body pile predominantly pale, except few black pile

136 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

on vertex and scutum; basoflagellomere elongated (1.8 times as long as wide) obviously
concave dorsally, arista 1.8 times as long as basoflagellomere (Fig. 11G—I); tergum 2
with pale lateral maculae; metafemur incrassate with medium long pile on ventral sur-
face, length approximately one third of its width (Fig. 8D); male genitalia: posterior
surstyle lobe with lateral hump; apical part of anterior surstyle lobe rhomboid; lingula
medium size (Fig. 9G—I). Related to Merodon serrulatus from which differs in absence
of medial fossette (Fig. 11H), present in geographically related Iberian populations
of M. serrulatus (Fig. 3B), molecular data and distribution (Fig. 7). Related to M@.
bequaerti, but differs by shorter pile on ventral margin of metafemur in both sexes
(Fig. 8D), narrower and oval to triangular apical part of anterior surstyle lobe (Fig.
9G), with rounded margin in MZ. bequaerti (Fig. 9A), and light yellow and less dense
marginal pile on apical part of anterior surstyle lobe (Fig. 9G, K: al), dark brown and
dense in M. bequaerti (Fig. 9A: al, J).

Redescription (based on the material from type locality, Middle Atlas, Azrou).
Male. Head. Antennae black to dark brown; basoflagellomere elongated, ca. 1.8 times
as long as wide, and ca. 2.5 times as long as pedicel, concave dorsally with acute apex;
fossette dorsolateral; arista dark and thickened at basal one third, covered with dense
microtrichia, ca. 1.8 times as long as basoflagellomere (Fig. 11G, H); face and frons
black with gray microtrichia, face covered with dense whitish, and frons with yellowish
gray pile; oral margin shiny, with small lateral microtrichose area; lunule shiny black,
bare; vertex shiny black, except in front of anterior ocellus, covered with microtrichia;
vertex isosceles, with long, pale whitish yellow pile, mixed with black pile on the ocellar
triangle; ocellar triangle isosceles; eyes covered with dense pile; occiput with gray-yel-
low pile, ventrally covered with a dense, gray microtrichia; eye contiguity 8—11 facets
long; vertical triangle: eye contiguity: frons = 3: 1 : 3.

Thorax. Scutum and scutellum black with bronze luster, covered with dense,
erect, usually yellow pile; sides of scutum at wing basis with patch of black pile or fas-
cia of short black pile and few black pile between wing basis; scutum with two micro-
trichose vittae, anteriorly connected and posteriorly reaching the scutellum; anterior
half of scutum dull; posterodorsal part of anterior anepisternum, posterior anepis-
ternum (except anteroventral angle), anterior anepimeron, dorsomedial anepimeron,
and posterodorsal and anteroventral parts of katepisternum with long, dense pale
yellow pile and grayish microtrichia; wings entirely covered with microtrichia; wing
veins brown; calypteres and halteres pale yellow; legs mostly black, except brown
tarsi ventrally in some specimens; pile on legs pale yellow; metafemur moderately
incrassate, ca. three times longer than wide; pile on postero- and anteroventral surface
medium long, and ca. as one third of width of metafemur, approximately the same
length as pile on dorsal surface (Fig. 8D).

Abdomen. Tapering, 1.2 times longer than mesonotum; terga dark, except for a
pair of pale yellow-orange, triangular, lateral maculae on tergum 2; terga 3 and 4 each
with a pair of white microtrichose and oblique fasciae (on tergum 2 triangular); pile on
terga all yellow; sterna dark brown, covered with long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, ca. 1.5 times
longer than wide, covered with dense, short pile (Fig. 9G, J: al); posterior surstyle lobe

Revision of the Merodon serrulatus group 137

oval with basolateral protrusion (lateral hump) (Fig. 9G, H: bp); hypandrium sickle-
shaped, without lateral projections; lingula medium size (Fig. 9I: 1).

Female. Similar to the male except for normal sexual dimorphism and for the fol-
lowing characteristics: antennae with rounded tip, basoflagellomere ca. two times long-
er than wide (Fig. 111); frons with broad microtrichose vittae along eye margins; frons
covered with variable pilosity, from mostly gray-yellow until predominantly black;
ocellar triangle covered with black pile; lateral side of terga, anterior two thirds of ter-
gum 2 and all of tergum 5 with yellow pile; terga 2—4 with short adpressed black pile.

Distribution. Merodon sophron is distributed in north-western Africa (Morocco)
(Fig. 7).

Ecology. Preferred environment: forest/open ground; open areas in evergreen oak
maquis, dry Pinus forest; unimproved grassland and tracksides (Fig. 35C). Flowers
visited: no data. Flight period: May-June.

Type material. Holotype [original designation by Hurkmans (1993: 168)]. Morocco
¢ 4; Azrou; 33°25'00"N, 5°20'00"W; 29 May 1925; E. Hartert leg.; NHMUK (studied).

Other material. Morocco * 1 4; Azrou; 30°40'00"N, 7°30'00"W; 31 May 1953;
G. L. Spoek leg.; NBCN ¢ 2 So; Moyen Atlas, Azrou; 30°40'00"N, 7°30'00"W; 19
Jun. 1928; R. Benoist leg.; MNHN PM0344, PM0350 ¢ 1 2; Moyen Atlas, Azrou;
30°40'00"N, 7°30'00"W; 16 Jun. 1928; R. Benoist leg.; MNHN PM0371 ¢ 1 9;
Middle Atlas, Azrou; 33°24'51"N, 5°11'36"W; 1789 m a.s.l.; 25-26 Jun. 2014; A.
Vujic, S. Radenkovié, J. Acanski, S. Veseli¢ leg.; FSUNS 07044 ¢ 1 3; Moyen Atlas,
Azrou; 33°25'48"N, 5°12'36"W; 16 Jun. 1928, R. Benoist leg; MNHN 22624 ¢ 1 3;
Middle Atlas, Maknes, Azrou; 33°25'48"N 5°12'36"W; 1800 m a.s.L.; 25 May 1995;
C. Kassebeer leg.; FSUNS 02496.

Merodon trianguloculus Vujié, Likov & Radenkovi¢ sp. nov.
http://zoobank.org/343 FE864-04B3-4B5A-BC75-217 DDEFA7CDE
Figs 23A—D, 33A—C, 34A—C

Diagnosis. Medium sized (7.5—11.6 mm), dark brown species with characteristic large
silver microtrichose fasciae on terga 2—4 in males (Fig. 23C), and silver michrotrichose
ornamentation on scutum in both sexes (Fig. 23A, B); basoflagellomere with rounded
apex, 1.6—1.8 times longer than wide in male (Fig. 33A, B).

Description. Male. Head. Antennae black to dark brown; basoflagellomere
rounded, 1.6—1.8 times as long as wide, and ca. 2.3 times as long as pedicel; large
fossette dorsomedial and dorsolateral; arista brown and thickened at basal one third,
covered with dense microtrichia, ca. 1.8 times as long as basoflagellomere (Fig. 33A,
B); face and frons black with gray microtrichia, face covered with dense whitish gray,
and frons with yellowish gray pile; oral margin shiny with microtrichose lateral areas;
lunule shiny black, bare; vertex covered with gray microtrichia; vertex isosceles, with
long, pale whitish yellow pile mixed with black pile on the ocellar triangle; ocellar tri-
angle equilateral; eyes covered with dense pile; occiput with gray-yellow pile, covered
with a dense, gray microtrichia; eye contiguity 8—12 facets long.

138 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Figure 34. Male genitalia. A Merodon trianguloculus sp. nov., epandrium, lateral view B Merodon trian-
guloculus sp. nov., epandrium, ventral view C Merodon trianguloculus sp. nov., hypandrium, lateral view.

Abbreviations: al—anterior surstyle lobe, bp—basolateral protrusion, |-lingula. Scale bar: 0.2 mm.

Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect,
yellow pile; scutum with conspicuous silver microtrichose ornamentation (Fig. 23A);
posterodorsal part of anterior anepisternum, posterior anepisternum (except anter-
oventral angle), anterior anepimeron, dorsomedial anepimeron, and posterodorsal and
anteroventral parts of katepisternum with long, pale yellow pile and grayish micro-
trichia; wings entirely covered with microtrichia; wing veins brown; calypteres and
halteres pale yellow; legs mostly black, except yellowish tip of femora, basal and apical
part of tibiae and brown tarsi ventrally; pile on legs pale yellow; metafemur moderately
incrassate, ca. four times longer than wide; pile on metafemur long, and ca. half to two
thirds of width of metafemut.

Abdomen. Tapering, 1.2 times longer than mesonotum; terga dark, with broad
silver microtrichose fasciae; tergum 2 with pale orange lateral maculae; pile on terga all
yellow (Fig. 23C); sterna dark brown, covered with long whitish yellow pile.

Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, approximate-
ly as long as wide, covered with dense, short pile (Fig. 34A: al); posterior surstyle lobe
oval with basolateral protrusion (lateral hump) (Fig. 34A, B: bp); hypandrium sickle-
shaped, without lateral projections; lingula (Fig. 34C: 1).

Female. Similar to the male except for normal sexual dimorphism and for the
following characteristics: basoflagellomere ca. 1.8 times longer than wide, fossette dor-
solateral (Fig. 33C); frons with microtrichose vittae along eye margins; frons covered
with mostly gray-yellow pile mixed with black ones; ocellar triangle covered with black
pile; ventral margin of metafemur with sparse pilosity, only individual pile longer;
lateral side of terga, anterior two third of tergum 2 and all tergum 5 with whitish pile;
terga 2—4 with short adpressed black pile medially; microtrichose fasciae on terga 3 and
4 narrower (Fig. 23D).

Etymology. The name trianguloculus derives from the Latin adjective triangulus
(triangular) and Latin noun Joculus (spot) and describes the distinctive triangular silver
pollinose fasciae on the abdomen.

Revision of the Merodon serrulatus group 139

Figure 35. Different types of habitats of Merodon serrulatus species group. A Lesvos (Greece), habitat of
Merodon opacus sp. nov., Photograph by Ante Vuji¢ B Siberia, Teletskoye Lake (Russia), habitat of Mero-
don serrulatus, Photograph by Jeroen van Steenis C Morocco, habitat of Merodon sophron, Photograph
by Ante Vuji¢ D Tajikistan, habitat of Merodon nigrocapillatus sp. nov., Photograph by Anatolij Barkalov.

Distribution. Merodon trianguloculus sp. nov. was recorded only in Turkmenistan
(Fig. 7).

Ecology. Preferred environment: open areas extending to the forest zone; unim-
proved grassland; adults resting on the stones and in flight between grasses at the top
of Dushak Mountain. Flowers visited: no data. Flight period: May-June.

Type material. Holotype. TuRKMENISTAN * 3; 120 km SW Geok-Tepe town;
38°10'31"N, 57°58'01"E; 11 May 1988; A. Barkalov leg.; SZMN 05818. Origi-
nal label: “HOLOTYPE of Merodon / trianguloculus Vuji¢, Likov / et Radenkovi¢
sp.n. 2019” [red label], “T'ypxmenua, 120 km / 103 Veox—Teme 11.Y. 1988 / C6.A.
Bapxaaos”, “05818” (See Supplementary file 5: Figure 5). Paratypes. TURKMENISTAN
¢1 2; 15kmk-s pos. Firjuza settlement, Dushak Mountain; 18 May 1988; A. Barka-
lov leg.; SZMN 05819 © 1 2; same data as for preceding; SZMN 05837 ¢ 1 9; Centr.
Kopetdag g. Dusak; 2100-2300 m a.s.l.; 6 Jun. 1986; Dubatolov leg.; SZMN 05844
13; Firjuza settlement 15 km zap., Dushak Mountain; 16 May 1988; A. Barkalov leg.;
SZMN 05816 © 1 4; same data as for preceding; 8 May 1987; SZMN 05840 ¢ 1 3;
120 km SW Geok-Tepe town; 38°10'31"N, 57°58'01"E; 11 May 1988; A. Barkalov
leg.; SZMN 05817.

140 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Merodon trizonus (Szilady, 1940) nomen dubium

Remarks. The identity of Merodon trizonus remains unclear. The species was described
based on two male and two female syntypes labelled “La Calle [el Kala], Algeria” and
“Ain Draham, Tunisia’, which were not examined. Originally, the syntypes were lo-
cated in the Hungarian National Museum in Budapest, but the Diptera collection was
destroyed by a fire in 1956. The description of Szilady (1940) is incomplete and based
on a few differences from the related species M. hirsutus. The types of M. trizonus are
assumed lost and the description is insufficiently accurate to associate name to one of
these species. Currently, two species from the M. serrulatus species group, to which M.
hirsutus belongs, occur in northern Africa, namely M. bequaerti and M. sophron. There-
fore, we propose to leave the name Merodon trizonus (Szilady, 1940) as nomen dubium.

Key to the Merodon serrulatus species group

1 Posterior part of mid coxa without long pile (Merodon avidus-nigritarsis

| FETY=2T cl) 8 ASE nnn BN a vn PENI ise RS MAP DEzE Vo SPEAR 2
— Posterior part of mid coxa with long pile 0.0.0.0... other Merodon lineages
2 Taxa with characteristic basolateral protrusion (lateral hump) on posterior

surstyle lobe (as on Figs 1, 6: bp). Species with dark scutum and usually
whitish microtrichose fasciae on terga 2—4, at least in females (as on Fig.
15C); tergum 2 usually with a pair of reddish orange lateral maculae; abdo-
men elongated, usually narrow and tapering, slightly longer than scutum and
scutellum together (as on Figs 26, 29); legs mostly black; metafemur incras-
sate (as on Fig. 4); tarsi black dorsally and dark brown ventrally; antennae
usually dark; basoflagellomere usually obviously concave dorsally (as on Figs
11, 12); male genitalia: apical part of anterior surstyle lobe more or less of
rhomboid to triangular shape, covered with dense short pile (as on Fig. 1:
al); cercus rectangular, without prominences (as on Fig. 14A: c); hypandrium
elongated and sickle-shaped (as on Fig. 1); lateral sclerite of aedeagus finger-
like with basal thorn-like process (Fig. 1G: s); lingula usually present (as on
Fig. 1E: 1) Merodon serrulatus group) ............ccccscessceseeseeseeseesseeseeseeeenes 3
- Species without basolateral protrusion (lateral hump) on posterior surstyle lobe
in males and with different combinations of characters in females............cccceeee

..other species groups belonging to the Merodon avidus-nigritarsis lineage”

3 id Le tency eee CERISE va Se, SOLOS. R mt WN RTE Ome A nee RS Oe. Shies WER Mn Ce I 4
= Gala cd (oC RNRRS cen Re ee SN os SPP NRC eek SORE O ES REI EES oe RAE ee 5 19
4 Eyes dichopeicr as Ondrioss OB DVN \amsere ese ter. canton teu: Loar obteae te: 5
— EvecchOlOptct as On Pia ke aie soteussasasne toto ustedten MutadansGitteact uta lurataaaautuednts 6

* not treated here
** not treated here

10

1]

12

Revision of the Merodon serrulatus group 141

Black species with predominantly black body pile, especially on thorax (Fig.
26A, B); terga 2—4 bare or with indistinct microtrichose fasciae (Fig. 26A);
distribution: Tajikistan (Fig. 7)... Merodon nigrocapillatus sp. nov.
Species with olive-brown reflection, predominantly covered with pale yellow
pile; terga 2-4 with conspicuous lateral microtrichose fasciae (Fig. 15A); dis-
tribution: Kyrgyzstan and Kazakhstan (Fig. 7) .Merodon disjunctus sp. nov.
Bluish species (Fig. 29A) with dark macula on the medial part of the wing
(Fig. 29C) and whitish pale body pile; distribution: Uzbekistan (Fig. 7) .......

danas Point tins anes Poet th sells bycsed eth estoteeduaten estate Merodon nigropunctum sp. nov.

Dark brown species without dark macula on WiNgS........ cs eeeeseeseceeeeeeeeeee 7
Tergum 2 entirely dark brown to black (as on Fig. 21A) wo... cee eeeeeeeeeeees 8
Tergum 2 with yellow-orange lateral maculae (at least small ones) (as on Fig.
DS Fi) Se ee > Fee er Cee Ne PEN RenpRNe - Syne wes nerernty Sent 29 -Ge 2p-—eee s SEs 10

Scutum without black pile, except few black setae at wing basis in some speci-
mens; terga 2—4 with a conspicuous microtrichose fasciae (as on Fig. 21A)...9
Scutum with black pile, at least on fascia between wing basis; terga 2—4 with
a less conspicuous microtrichose fasciae (as on Fig. 2A) wo... ceeeeseeeeeeeeeeees
eevee peers tem, Merodon serrulatus (Wiedemann in Meigen, 1822) (part)
Dorsolateral pile on metafemur dense and longer (Fig. 13C); terga with longer
and erect pile (Fig. 21F); tergum 2 shiny; posterior surstyle lobe with big lat-
eral hump, clearly visible in ventral view (Fig. 14E, F: bp); distribution: Syria,
Israel and south-eastern Turkey (Fig. 7)........... Merodon hirsutus Sack, 1913
Dorsolateral pile on metafemur shorter (Fig. 13A); terga with shorter pile,
adpressed at tergum 4 (Fig. 21C); tergum 2 dull; posterior surstyle lobe with
small lateral hump, less distinct in ventral view (Fig. 14H, I: bp); distribu-
tion: Lesvos Island (Greece) and western Turkey (Fig. 7) ......ceseeseeeeseeseeeees
FEE NE eh ek TR ALS ne ANY SE Merodon opacus sp. nov.
Terga 3 and 4 with a pair of broad silver microtrichose maculae (Fig. 23C);
scutum with characteristic silver microtrichose ornamentation (Fig. 23A);
distribution: Turkmenistan (Fig. 7) ........... Merodon trianguloculus sp. nov.
Terga 3 and 4 without or with a less conspicuous pair of broad silver micro-
trichose tasciae{as-on Piet) OM sites eta cn uacs cadence ea Rognvon dean doncdanes Sareea 11
Terga 3 and 4 without microtrichose fasciae..........scseesseseeseceseeseeeeeeseeaeeeee 12
Terga 3 and 4 with a pair of white microtrichose, oblique fasciae (as on Fig.
S04 Up kanes dbtmnenty sala, / creel renin cbetils senha chee Rel, Pesmnlardgrert ie (PR Rea Marrs | 13
Metafemur strongly curved (Fig. 8C); body pile longer, clearly visible on ter-
eum 4 (Fig. 10D); distribution: Iberian Peninsula (Fig. 7)... seeeseeeeeeeees
ees eae ae Re En Eee ee oe Merodon sacki (Paramonov, 1936)
Metafemur less curved (Fig. 8A); body pile shorter, clearly visible on tergum
4 (Fig. 10B); distribution: north-west Africa (Fig. 7)......sesceeesseseeeeeeceeeeees
tet et ok oo te eee Merodon bequaerti Hurkmans, 1993 (part)

142

13

14

15

16

ii,

18

Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

Antennae reddish yellow; basoflagellomere short and broad, ca. 1.2 times as
long as wide, with large dorsal to dorsolateral fossette (Fig. 19E—G); tergum
2 with large reddish yellow lateral maculae (Fig. 23E); tergum 3 laterally red-
dish or brown; tibiae and tarsi partly reddish brown; metafemur incrassate
with long pilosity as long as half of width of metafemur (Fig. 22A); distribu-
tion: Japan and China (Fig. 7) ..... Merodon kawamurae Matsumura, 1916
Antennae dark brown/black; basoflagellomere elongated; legs mostly black .... 14
Abdomen broad, tergum 2 at least 2.5 times wider than long (as on Fig. 24A);
metatemurinerassate-and.curved fas: OnsFig FOG) ict. cvccrsnrosncthvceconnpencestvuece 15
Abdomen narrower, tergum 2 ca. two times wider than long (as on Fig. 2);
metafemur less incrassate and with almost straight lateral margin (as on

PLE ie letcecet tas ecteseeigest vain entangling ech eabhimncbitn cade reupltr bested etn etek enti iccuatvsntens 16
Pile on ventral margin of metafemur very short (Fig. 22C); distribution:
Grete Islands Greece) (Fie. F))fsssccseSi teases eeens Merodon medium sp. nov.

Pile on ventral margin of metafemur long and dense (Fig. 8A); distribution:
north-west Africa (Fig. 7)....... Merodon bequaerti Hurkmans, 1993 (part)
Male genitalia: basolateral protrusion (lateral hump) on posterior surstyle
lobe reduced (Fig. 14A: bp); distribution: western Turkey (Fig. 7) ............
Be Sh ED ch ud le ie RM Bette le eek OriesteiT ee Merodon defectus sp. nov.
Male genitalia: basolateral protrusion (lateral hump) on posterior surstyle
lobe well developed (as on Fig. 1, 6: bp), visible at least from ventral view
Gesteceal 225 Gis 0) 0) ener Ren oe Se RoR aE SR ee 17
Medial fossette-alnserity israel Tel) hcl shane cautastlog cour tesvesttea chaleencsioeutct 18
Medialtrossette*preseiit (F115 B)tirk ists. Sst oseeB eh ome venutro Soma cohol eS caiai,
Pe, eae... wee Merodon serrulatus (Wiedemann in Meigen, 1822)°
GenBank acc. no. MN623564-MN623581; distribution: Palaearctic, ex-
tending from France in the west to Turkey in the south-east, and to Siberia
LOWARGHORLNE CASte UIC’. 72), Bastyr wheyrowmius vhstinavaes hau nteaV neal hor eea WL wwed tasawts Seawees
eaten Sete maul hae Merodon serrulatus (Wiedemann in Meigen, 1822) (part)
GenBank acc. no. MN623540; distribution: north-west Africa (Fig. 7) ........
eee ne ae Pete ee ee Merodon sophron Hurkmans, 1993
Teroum2-dark(as.on Fie. 26C 1) Sse weet ee 20°
Tergum 2 with reddish lateral maculae (as on Fig. 23F) oo. eeeeeeeeeeee 22
Black species (Fig. 26C, D); body covered with predominantly black pilosity,
especially on thorax; distribution: Tajikistan (Fig. 7) oe... ee eesesseeseeeeeseeeeeeees
prt Fie Shs eee eka gts oc hein shane Merodon nigrocapillatus sp. nov.
Species with olive-brown reflection, predominantly covered with pale pile....21
Pile on dorsolateral margin of metafemur long (Fig. 13D); distribution: Syria
and south-east Turkey (Fig. 7) ......sceeeeeseeeee Merodon hirsutus Sack, 1913
Pile on dorsolateral margin of metafemur short (Fig. 13B); distribution: Les-
vos Island (Greece) and western Turkey (Fig. 7) .... Merodon opacus sp. nov.

* part: Iberian populations

** unknown female of Merodon nigropunctum sp. nov. probably keys out here

22,

23

24

25

26

27

Revision of the Merodon serrulatus group 143

Metafemur with long pile on the entire surface of ventral margin (as on 22B)
(unknown female of Merodon sacki (Paramonov 1936), probably keys out
|S 2) SR en al pace nes rane REN nih iP Sa RA RRND xpi ean NE Nhs Dhiea t aah eae aR 23
Metafemur with mostly short pile on ventral margin (as on Fig. 4C)........ 27.
Scutum with characteristic silver microtrichose ornamentation (Fig. 23B);
terga 3 and 4 with broad lateral microtrichose fasciae; distribution: Turk-

MICO SLATIE HIS. 27 \cactornatted .Sotut Ratt Sao Muale Merodon trianguloculus sp. nov.
Scutum with indistinct microtrichose vittae; terga 3 and 4 with narrower
Figcieed Mactisleoiselelo’ehiom 7 h(c penance Soom nf Sea ee Ronin 24
Scutum at wing basis with only yellowish pilosity; metafemur with long,
devisedoisal piles OmgiGr a2 By as sustestunteeediabes tsa etens oun ieteatied 25
Scutum at wing basis with short black pile; metafemur with sparse dorsal pile
(aston hig: Oi) hm ser teen teen: emer a sea rosea ase een dela eves ntact nemo 26

Blackish species; terga 3 and 4 with broad lateral microtrichose fasciae; baso-
flagellomere dark brown to black; distribution: Kyrgyzstan and Kazakhstan
CVC a7) cues Meaierevh Sento g ou inever air enanelercsnngegacesuaces3 Merodon disjunctus sp. nov.
Brownish species; terga 3 and 4 with narrower lateral microtrichose fas-
ciae (Fig. 23F); basoflagellomere yellowish; distribution: Japan and China
(PIG a) orient Wraawecbiaccrstagtrsta ts Merodon kawamurae Matsumura, 1916
Distributions nore west 7uricd, (FIG 7). | Gea setnccstnes tadhtn anlteessr tat sdneeasgniden
bdr hae, Seer Rl A NP a Asp, Rn hae Merodon bequaerti Hurkmans ,1993

Aiud |e ai T, FL Merodon serrulatus (Wiedemann in Meigen, 1822) (part)
Females of these three species can be separated by distribution and genetic data:
GenBank acc. no. MN623540. Distribution: North-west Africa (Fig. 7) ......
o Riodiai gaan aS eat oe ea ean hres esee Merodon sophron Hurkmans, 1993
GenBank acc. no. MN623564-MN623581. Distribution: Palaearctic, ex-
tending from Iberian Peninsula in the west to Turkey in the south-east and
toward Siberia in the noktnreast, (Pies) Jisse Masel ttseadeaaal ccs Aa dueasdecs a tetldesine
Linh Renee ee rd Merodon serrulatus (Wiedemann in Meigen, 1822) (part)
Distribution: western Turkey (Fig. 7)... eee Merodon defectus sp. nov.

Molecular inference

The final aligned and pruned dataset including two-gene data matrix (COI+28S rRNA)

comprised 1,859 nucleotide characters (421 parsimony informative sites) pertaining to

81 specimens (79 in-group specimens of the studied genus Merodon lineages along
with two outgroups). The final number of aligned sites for COI gene (concatenated 3’
and 5’ fragments of the gene) included 1,273 nucleotides, while 586 nucleotide charac-
ters (with gaps) were included in analyses for the D2-3 region of the 28S rRNA gene.

144 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

AU739 Platynochaetus maquarti
COOOCROO OCMC OCPROOO CHROMO EU13 Eumerus grandis:
weris | Af nalans lineage

OOOCOOCOMHEOO— N20 M.nanus
AU760 M.chalybeus

OK OCOCCAHROCOOBOCO— Y665 M.aureus

84 cocco gE EENTN GG 00800090G0G0003000RCODODOO CORE CO HOOIMBOOS 393 M.cinereus

CODOCCOCODOGODOSO- DPCOCOCOSOOPOPOCOOOS! Y175 M.cabanerensis.

64 OOO OPOCOHEOCOCOCOMOODOCOCOODODOSOCOOCOMEBOOS 2078 M.desuturinus
OO

Sonesoceds:

M. aureus lineage

tris
oeCOORO— Y706 M.albifrons M. albifrons+desuturinus lineage

COCO en OOOO COCO OOO OOOO OHARTION ¥392 M.constans
HOBOOCOOOCODOCOOPOCOMPOHHMEDS 2077 M.ruficornis
56 OS: o— $90 M.avidus
OOOOCIC 95 OnmoeOCSCS0C0m Oo- Y700 M.testaceus
Pe ooocoe ef Oo O— ¥1272 M.alagoezicus
OR OCOOCODOCOOCECOCOROCOOO GOOG 870 M.nigritarsis
on > OOSOC00Oe= 11278 M.erivanicus

B4
OO

oO
OOO COCOOOMOMOCODODODOOO

C0000

HECCOGOBOOO ¥1183 M.aurifer
OOOCOCOOSO00CO— 11247 M.clunipes:
748 M.clavipes
OO- Y704 M.velox
OOOOCODOOODOOOIOOSOCOGOOOGOOCOOHMROEOOOBOTOD 717 M.fulcratus
SXOONOOOOSIOOCOOCOSOGOIOCO oCococHe= 2083 M.ottomanus
00 —o— TS548 M.nigrocapillatus sp.n. Tajikistan
$649 M.nigrocapillatus sp.n. Tajikistan
AU295 M.medium sp.n. Greece,Crete
100 AU296 M.medium sp.n. Greece,Crete
54 COMODHOOCCCCODO#OOO—O— AUZ97 M.medium sp.n. Greece,Crete
OOOCO00000- 000— AU298 M.medium sp.n. Greece,Crete
AU299 M.medium sp.n. Greece,Crete

97 AU340 M.trizonus Morocco Il
6-— MS14 M.bequaerti Morocco .
96 97 MS15 M.bequaerti Morocco M. Bere
OOOCOCOODO 84-— MS$12 M.bequaerti Morocco ngiarsis lineage
MS13 M.bequaerti Morocco

r— AU300 M.defectus sp.n. Turkey,Bozdag

DBOCOCOCOOOOCOCOCOOOOOOCOK

COCO 00000 0000-

XIII MEOH IOI OI IH OI

AU301 M.defectus sp.n. Turkey,Bozdag

AU302 M.defectus sp.n. Turkey,Bozdag

AU303 M.defectus sp.n. Turkey,Bozdag
8 Sp.n.

00000
rkey,Bozdag

AU309 M.opacus sp.n. Turkey,Bozdag M. serrulatus
100 TS587 M.opacus sp.n. Greece,Lesvos group
Y701 M.opacus sp.n. Greece,Lesvos
TSS588 M.opacus sp.n. Greece,Lesvos
100 _ 4U778 M.sacki Spain,La Corte
AU779 M.sacki Spain,La Corte

70 MS3 M.serrulatus Russia, Altai
foes M.serrulatus Russia, Altai
MS5 M.serrulatus Russia, Altai

r— MS6 M.serrulatus Greece,Olympus
871 ws7 M.serrulatus Greece,Olympus.
MS8 M.serrulatus Greece, Olympus
“— MS9 M.serrulatus Greece,Olympus

AU292 M.serrulatus Greece,Peloponnese
63 AU293 M.serrulatus Greece,Peloponnese
7 AU290 M.serrulatus Greece,Peloponnese
AU291 M.serrulatus Greece,Peloponnese

AU294 M.serrulatus Greece,Peloponnese
p-O- AUS40 M.defectus sp.n. Turkey,Isparta
gol AU941 M.defectus sp.n. Turkey,lsparta
Lod} — auas? M.defectus sp.n. Turkey,|sparta
}— AU944 M.defectus sp.n. Turkey,lsparta
oe AU945 M.defectus sp.n. Turkey,Isparta

AUT77 M.serrulatus Spain,La Corte
64 AU341 M.serrulatus Spain,Sierra Nevada
AU342 M.serrulatus Spain,Sierra Nevada
AU343 M.serrulatus Spain,Sierra Nevada
AU344 M.serrulatus Spain,Sierra Nevada
AU345 M.serrulatus Spain,Sierra Nevada

Figure 36. Strict consensus tree of 41 most parsimonious trees from the analysis of combined COI

mitochondrial and 28S nuclear genes sequences. Length 2093 steps, Consistency Index (CI) 37, Reten-
tion Index (RI) 65. Bootstrap support values are depicted near nodes (2 50). Filled circles represent non-
homoplasious changes and open circles are homoplasious changes. Four lineages observed in the genus

Merodon, as well as the M. serrulatus group, are marked on the tree.

Both obtained phylogenetic trees (Maximum Parsimony, Fig. 36 and Maximum
Likelihood, Supplementary file 7: Figure 7) resolved the four previously described
lineages as clades, while the M. serrulatus species group was recovered as monophy-
letic within the Merodon avidus-nigritarsis lineage (MP = 54, ML = 75). Within
the serrulatus species group, specimens belonging to M. nigrocapillatus sp. nov., M.
medium sp. nov., M. bequaerti and M. sacki were clearly grouped together with high
bootstrap support (MP = 100, ML = 100; MP = 100, ML = 100; MP = 97, ML = 94;
MP = 100, ML = 100, respectively). The single sequenced specimen of M. sophron
was resolved as sister taxon of M. bequaerti. Unfortunately, although morphologi-
cally differentiated, specimens identified as M. defectus sp. nov. clustered with M. ser-
rulatus in a clade without support, but also with MM. opacus sp. nov. in another clade
without support. These three species together with M. sacki were resolved in a group
with high support value. High level of inter-population molecular variability within
M. serrulatus species was also detected.

Revision of the Merodon serrulatus group 145

Discussion

Taxon delimitation and integrative taxonomy

Likov et al. (2019) reported the monophyly of the Merodon avidus-nigritarsis lineage.
Within this lineage, the Merodon serrulatus species group is supported in our phyloge-
netic analyses.

The M. serrulatus species group comprises six already described species (Merodon
bequaerti, M. hirsutus, M. kawamurae, M. sacki, M. serrulatus, and M. sophron) and
seven new species described here. Based on the present results, six species of this group,
namely M. disjunctus sp. nov., M. kawamurae, M. medium sp. nov., M. nigrocapillatus
sp. nov., M. nigropunctum sp. nov., and M. trianguloculus sp. nov., are delimited on dif-
ferences of morphological characters. Moreover, two pairs of very similar species can be
separated from other species of the group by some distinct characters, but the distinc-
tion between the species in each pair is based on characters with more subtle differences.
These two pairs are M0. bequaerti | M. sacki, with the metafemur incrassate and long pile
on postero- and anteroventral surface of the metafemur, and M. hirsutus | M. opacus sp.
noy., with tergum 2 dark, without yellow-orange lateral maculae in both sexes.

The remaining three species within the M. serrulatus species group are morphologi-
cally very similar to each other. Merodon defectus sp. nov. has subtle, but stable differ-
ences in structures of the male genitalia serving as diagnostic characters (lateral hump
on posterior surstyle lobe reduced). Closely related and very similar, M. serrulatus and
M. sophron are distinguished by molecular data, in addition to a clear morphological
diagnostic character in males (presence or absence of medial antennal fossette).

Using different methodologies to assess various aspects of the diversity of the
genus Merodon, previous authors (Mengual et al. 2006; Marcos-Garcia et al. 2007,
2011; Stahls et al. 2009; Radenkovié et al. 2011; Vujié et al. 2012, 2013, 2015) have
shown the potential of the integrative taxonomy to indicate cryptic taxa, to define new
species and to point out different evolutionary lineages. The integration of multiple
data sources, combining different molecular (Popovi¢ et al. 2014, 2015; Acanski et al.
2016; Sasié et al. 2016; Kodis Tubié et al. 2018; Radenkovié et al. 2018a), morphologi-
cal (Popovicé et al. 2015), distributional (Radenkovié et al. 2018a), and environmental
(Popovié et al. 2015; Vujié et al. 2015; Sadié et al. 2016) information, has proven to
be significant in re-evaluating taxonomic delimitations within the Merodon genus. Al-
though results of this integrative approach have not been always congruent (Mengual
et al. 2006; Stahls et al. 2009; Popovicé et al. 2015; Radenkovi¢ et al. 2018a).

In the present study we applied this integrative approach, i.e., to combine mor-
phology, genetic data, and distribution, to support the taxonomic status and systematic
decisions made for the . serrulatus species group. For example, the species M. sophron
and M. serrulatus, although morphologically similar, are conspicuously separated from
each other based on molecular data. The same situation is found between M. bequaerti
and M. sacki. In contrast, the morphologically distinct species M. defectus sp. nov., M.
serrulatus, and M. opacus sp. nov. cluster together in the molecular analysis. Discord-

146 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

ance between morphological and molecular data has been observed in some previous
studies concerning closely related taxa within the family Syrphidae (e.g., Stahls et al.
2009; Francuski et al. 2014; Haarto and Stahls 2014), as well as in recently conducted
studies on Merodon species groups (Likov et al. 2019). In the present study the mo-
lecular data for M. defectus sp. nov. show some interpopulation differentiation: while
the specimens from Bozdag (Turkey) were resolved in the same cluster with MZ. opacus
sp. nov., the specimens of M. defectus sp. nov. from Isparta (Turkey) and M. serrulatus
were resolved in another cluster. Unfortunately, these two clades do not have sup-
port and the whole cluster, including M. sacki, could be resolved in a large polytomy
when collapsing nodes without high support. Different molecular profile of different
populations of one species was also detected by Likov et al. (2019). The suggested rea-
sons for the low COI divergence between these species are retained polymorphism or
mitochondrial introgression between the taxa, as it has been hypothesized in previous
studies (e.g., Stahls et al. 2009; Francuski et al. 2014; Haarto and Stahls 2014).

It is important to do further taxonomic research with the populations of Merodon
serrulatus with high inter-population morphological and genetic variability. These pop-
ulations may be also geographically isolated and are posited to exhibit low genetic flow.
The very wide distributional range of M. serrulatus, extending from Iberian Peninsula to
Mongolia, is highly unusual in the genus Merodon, thus exemplifying a complex popu-
lation structure that might contain evolutionary units at different levels of speciation.

Immature stages

One of the main reasons for the gap in extant knowledge on the immature stages of
Merodon species is the difficulty of finding specimens in the field, since host plants,
the larval food-plants and the breeding and oviposition sites, have not been recorded
for the great majority of Merodon species (Hurkmans 1993; Rotheray 1993; Speight
2018). The description of the puparium of Merodon opacus sp. nov. in this work is
based on a single specimen reared from the larva found in the soil near the bulbs of Fr-
tillaria, Gagea, Muscari, and Ornithogalum. \n extant studies, the immature stages of
Merodon species were linked to bulbous geophytes, mostly belonging to plant families
Asparagaceae (Ricarte et al. 2008; Andri¢ et al. 2014; Preradovi¢ et al. 2018), Iridaceae
(Stuckenberg 1956) and Amaryllidaceae (Heiss 1938; Ricarte et al. 2017).

The morphology of the puparium of MZ. opacus sp. nov. shows similarities with the
puparium of M. avidus in terms of the morphology of the posterior respiratory process
(prp) and ornamentation of pupal spiracles (Preradovié et al. 2018). In fact, these spe-
cies share the button-shaped prp and the poorly defined outline of the spiracular plate,
whereas the spiracular openings of MM. opacus sp. nov. are less convoluted than those in
M. avidus. The pupal spiracles are stout in shape (almost as long as broad) and are clearly
shorter than in M. avidus, but share the reticulated ornamentation (polygonal pattern).

A single larva of Merodon opacus sp. nov. was found in the ground surrounded
with bulbs of different plant genera (Fritillaria, Gagea, Muscari, Ornithogalum). Recent

Revision of the Merodon serrulatus group 147

larval records suggest that groups of related Merodon species could have the same plant
genus as a host. These close relationships could be suspected between: MZ. constans
species group and Galanthus L. (Amaryllidaceae) [Popov and Mishustin (pers. comm)
confirmed that eight species of the constans species group feed on bulbs of eleven snow-
drop species], /Z. aureus species group and Crocus L. (Iridaceae) [Speight 2018; Popov
pers. comm.], and M. geniculatus species group and Narcissus L. (Amaryllidaceae) [i.e.,
M. eques (Fabricius, 1805) (see Pehlivan and Akbulut 1991), M. geniculatus Strobl,
1909 (see Ricarte et al. 2017), and M. neofasciatus Stahls & Vuji¢, 2018 (see Vuji¢ et
al. 2018)]. Based on these findings, we suggest that the host plant for the members
of the M. serrulatus species group should be a plant genus present on its large range,
extending from North Africa, throughout the entire Palaearctic region to Japan. Two
bulb genera with native ranges (WCSP 2019) fitting this distribution, Gagea and Fri-
tillaria (Liliaceae), might be the larval food-plants. Future research in this field could
thus focus on more detailed field work in areas characterized by numerous populations
of species from the M. serrulatus species group.

Distribution and species diversity

Being distributed from the Iberian Peninsula in the south-west, along the Mediterra-
nean and Balkan Peninsula, through Turkey and southern Russia to Siberia and Mon-
golia in the north-east, Merodon serrulatus is the species of the genus Merodon with
the largest distributional range. Other species of the M. serrulatus species group can
be found at the edges of this distributional range, albeit with a much more restricted
distribution. For example, /. sacki has been found in southern Spain, VM. medium sp.
nov. is endemic to Crete Island, whereas M. defectus sp. nov. and M. opacus sp. nov.
have been recorded in western Turkey, with the latter species also being found on
Lesvos Island, and M. Airsutus found in south-eastern Turkey, Israel and Syria. The
serrulatus species group includes two North-African species, i.e., MZ. sophron restricted
to Morocco, and M. bequaerti more widely distributed along the Mediterranean coast
of the African continent. Only one species of the group, M@. kawamurae, is found in
the Far East of the Palearctic region, i.e., in central and south-eastern China and Ja-
pan. It is worth noting that four of the seven newly described species are distributed
in Central Asia, the central and somewhat isolated part of the distribution range of
the M. serrulatus species group. Merodon disjunctus sp. nov. is found in Kyrgyzstan
and Kazakhstan, M/. nigrocapillatus sp. nov. has been collected in Tajikistan, whereas
M. nigropunctum sp. nov. and M. trianguloculus sp. nov. are found in Uzbekistan and
Turkmenistan, respectively.

The genus Merodon is known to be widespread in regions such as the Mediterranean
Basin, with high diversity of geophytes, whereby underground storage organs serve as
larval food sources for Merodon species (Ricarte et al. 2008, 2017). Such potential for the
development of a high diversity of Merodon taxa might explain their current geographi-
cal distributions (Vuji¢ et al. 2011, 2013). The highest number of Merodon species and

148 Ante Vuji¢ et al. / ZooKeys 909: 79-158 (2020)

the greatest endemicity level in the Mediterranean Basin was noted for the Anatolian
region (Vuji¢ et al. 2011), which represents the main center of Merodon diversity within
the Palaearctic region, along with the Iberian Peninsula (Marcos-Garcia et al. 2007).
The high number of endemic species in the eastern Mediterranean Basin has been sug-
gested to be related to the intense orogenic activity favoring isolation and allopatric
speciation (Vuji¢ et al. 2011). The biologically diverse Anatolian region, characterized
by a rich geological history, comprises of an extensive system of high mountain chains
and closed basins, thus providing a wide range of habitats. Throughout history, differ-
ent parts of this topographically complex area, connecting diverse geographic regions of
Asia and Europe, have served not only as natural barriers but also as highly important
refugia and corridors providing passages for species spreading (Vuji¢ et al. 2013, 2015).

Central Asia is characterized by many mountains exceeding 6,500 m in elevation, as
well as by major desert basins, which have thus far remained understudied. ‘This is particu-
larly the case for the alpine areas, and especially in terms of the invertebrate fauna (CEPF
2017). The very diverse flora of this region harbors a large number of endemics, including
many bulbous plants (CEPF 2017) which can support high diversity of Merodon taxa, in-
cluding the four endemic species of the M. serrulatus species group described here. Major
mountain ranges located in Central Asia represent an extensive zone for faunistic evolu-
tion and differentiation, not only ecologically, but also orographically and biogeographi-
cally (Mani 1968). Heterogeneous topography with various isolated habitats along altitu-
dinal gradients fosters high rates of speciation, species diversity and endemism. Climatic
fluctuations and tectonic processes throughout the complex geological history of this
region have contributed to its unique climate and have promoted high levels of floristic
diversification and alpine endemism, while also affecting the distributions and structure
of many taxa (e.g., Djamali et al. 2012; Zinenko et al. 2015). Having a long history as the
crossroads between east and west, this region has historically been subjected to high levels
of anthropogenic disturbance that continue to the present day, and populations of many
species have declined due to habitat modifications (Djamali et al. 2012; CEPF 2017). The
results yielded by the present study confirm previous conclusions emphasizing the impor-
tance of such underexplored regions as centers of endemicity, hosting habitats potentially

harboring hidden diversity within the genus Merodon (Vuji¢ et al. 2019).

Acknowledgments

We thank the curators of the museums listed in the Materials and methods for facili-
tating visits and loans for the study of specimens in their care. We are also indebted
to our professional English language editor for linguistic revision and editing of the
manuscript. This work was funded by the Ministry of Education, Science and Techno-
logical Development of the Republic of Serbia, Grant Nos. O1173002 and 11143002,
the Provincial Secretariat for Science and Technological Development (0601-504/3),
the H2020 Project “ANTARES” (664387) and the Scientific and Technological Re-
search Council of Turkey (TUBITAK, project number: 2130243). The work of A.V.

Revision of the Merodon serrulatus group 149

Barkalov was supported by the Russian Foundation for Basic Research. The authors
confirm that no competing interests exist.

Permission to collect biological specimens in protected areas was provided by the
competent authorities. The Greek material was collected under a permit issued by
Greek Ministry of Environment, Energy and Climate change (130276/1222), in Tur-
key by TUBITAK, and in Spain with permission N. Ref.: ENSN/FJSG/IMJ (232)
(Junta de Andalucia, Consejeria de Medio Ambiente y Ordenacién del Territorio).

Roles of authors: AV, LL, SRad, CPB, AB, RH, GS and SRojo performed the sam-
pling; AV, LL, SRad, NKT and MD conceived and designed the study; AV, LL, SRad,
NKT, MD, CPB, SR, AA, GS performed the experimental analysis, while AV, LL, SRad,
NKT, MD, AS, CPB, AB, RH, SRojo, AA, GS participated in data analyses. All authors
took part in draft preparation, contributed to discussions during preparation of the
paper, as well as read, commented on, and approved the final version of the manuscript.

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Supplementary material |

Figure S1

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovic, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: type specimens’ data

Explanation note: A Merodon bequaerti, holotype and labels B Merodon nigrocapillatus
sp. nov., holotype and labels.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl1

Supplementary material 2

Figure S2

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovic, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: type specimens’ data

Explanation note: A Merodon defectus sp. nov., holotype and labels B Merodon disjunc-
tus sp. nov., holotype and labels.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl2

156 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)

Supplementary material 3

Figure $3. Merodon medium sp. nov., holotype and labels

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovic, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: type specimens’ data

Explanation note: Merodon medium sp. nov., holotype and labels.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.supp13

Supplementary material 4

Figure S4

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovi¢, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: type specimens’ data

Explanation note: A Merodon nigropunctum sp. nov., holotype and labels B Merodon
opacus sp. nov., holotype and labels

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl4

Revision of the Merodon serrulatus group 157
Supplementary material 5

Figure $5. Merodon trianguloculus sp. nov., holotype and labels

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovi¢, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: type specimens’ data

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl5

Supplementary material 6

Figure S6

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovi¢, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: type specimens’ data

Explanation note: A Merodon tener, lectotype and labels B Merodon sacki, holotype
and labels.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl6

158 Ante Vujié et al. / ZooKeys 909: 79-158 (2020)
Supplementary material 7

Figure S7

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovic, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: phylogenetic data

Explanation note: Maximum likelihood tree based on analysis of combined COI mi-
tochondrial and 28S nuclear genes sequences. Bootstrap support values of the main
clades of the analysed Merodon serrulatus group of species are depicted near nodes
(> 50).

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl7

Supplementary material 8

Table S1. Data for the specimens used in the molecular analysis, including Gen-

Bank accession numbers

Authors: Ante Vuji¢, Laura Likov, Snezana Radenkovic, Natasa Kocis Tubi¢, Mihajla

Djan, Anja Sebié, Celeste Pérez-Bafién, Anatolij Barkalov, Riistem Hayat, Santos Rojo,

Andrijana Andri¢, Gunilla Stahls

Data type: molecular specimens’ dataset

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.909.46838.suppl8