Research Article |
Corresponding author: Laura Likov ( laura.likov@dbe.uns.ac.rs ) Academic editor: Ximo Mengual
© 2020 Ante Vujić, Laura Likov, Snežana Radenković, Nataša Kočiš Tubić, Mihajla Djan, Anja Šebić, Celeste Pérez-Bañón, Anatolij Barkalov, Rüstem Hayat, Santos Rojo, Andrijana Andrić, Gunilla Ståhls.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Vujić A, Likov L, Radenković S, Kočiš Tubić N, Djan M, Šebić A, Pérez-Bañón C, Barkalov A, Hayat R, Rojo S, Andrić A, Ståhls G (2020) Revision of the Merodon serrulatus group (Diptera, Syrphidae). ZooKeys 909: 78-158. https://doi.org/10.3897/zookeys.909.46838
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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 characters 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 Vujić, Likov & Radenković sp. nov., M. disjunctus Vujić, Likov & Radenković sp. nov., M. medium Vujić, Likov & Radenković sp. nov., M. nigrocapillatus Vujić, Likov & Radenković sp. nov., M. nigropunctum Vujić, Likov & Radenković sp. nov., M. opacus Vujić, Likov & Radenković sp. nov., and M. trianguloculus Vujić, Likov & Radenković 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 available 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 (Szilády, 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.
28S rRNA, COI, immature stages, lectotype, morphology, new species, new synonyms, taxonomy
The phytophagous hoverfly genus Merodon Meigen, 1803 contains more than 160 species distributed across the Palaearctic and Afrotropical regions (
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 (
Most recent publications pertaining to the genus Merodon have focused on particular species groups, within which the authors delimited individual species (
The Merodon serrulatus species group includes taxa with a characteristic basolateral protrusion on the posterior surstyle lobe (Fig.
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.
Most of the recently collected specimens were sampled by sweep net. Further specimens of the Merodon serrulatus species group were sourced from collections deposited in museums and universities which are listed below. Consisted total of 1,083 specimens 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, Turkmenistan, 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 beginning of a material citation), number and sex of specimen(s), locality data, geographical 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 interpretations provided in square brackets, where applicable.
A. S. coll. Axel Ssymank collection, Achtberg, Germany (ssymanka@t-online.de)
CEUA Colección Entomológica de la Universidad de Alicante, Alicante, Spain
D. D. coll. Dieter Doczkal collection, Munich, Germany (dieter.doczkal@gmail.com)
EMIT Entomological Museum of Isparta, Isparta, Turkey
G. V. W. coll. Guy Van de Weyer collection, Reet (Rumst), Belgium (guido.vandeweyer@skynet.be)
J. T. S. coll. John T. Smit collection, Utrecht, the Netherlands (John.Smit@naturalis.nl)
J. v. S. coll. Jeroen van Steenis collection, Amersfoort, the Netherlands (jvansteenis1@gmail.com)
M. B. coll. Miroslav Barták collection, Prague, Czech Republic (bartak@af.czu.cz)
M. H. coll. Martin Hauser collection, Sacramento, USA (martin.hauser@cdfa.ca.gov)
MAegean The Melissotheque of the Aegean, University of the Aegean, Mytilene, Greece
S. K. coll. Sakari Kerppola collection, Helsinki, Finland (sakari.kerppola@helsinkinet.fi)
S. S. coll. Süleyman Sarıbıyık collection, Kastamonu, Turkey
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 (Szilády, 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 lunule 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 photographing, CombineZ software (
Terminology adopted in the morphological descriptions follows
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) (
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 this period. An area of ca. 3m2, where the presence of many bulb species and adults of Merodon were reported the previous year was selected. The whole area was excavated 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 specimen 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
The specimens subjected to molecular analysis are presented in Supplementary file 8: Table
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
Diagnosis. Member species of the Merodon serrulatus species group exhibit a distinctive and characteristic basolateral protrusion (lateral hump) on the posterior surstyle lobe (Fig.
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, l–lingula, pl–posterior surstyle lobe, s–lateral sclerite of aedeagus. Scale bar: 0.2 mm.
Basoflagellomere 1.5–2.2 times as long as wide, usually obviously concave dorsally (as in Fig.
Merodon serrulatus antenna. A outer side, lateral view, male (Spain) B inner side, lateral view, 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). Abbreviation: f–fossette. Scale bar: 1 mm.
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.
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., M. nigrocapillatus sp. nov., M. nigropunctum sp. nov., M. opacus sp. nov., M. sacki, M. serrulatus, M. sophron Hurkmans, 1993, and M. trianguloculus sp. nov.
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 ventral margin; the longest pile as long as one third to half of width of metafemur (Fig.
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 medium sp. nov. G, H margin of anterior surstyle lobe marked with arrow. Abbreviation: bp–basolateral protrusion. Scale bar: 0.2 mm.
Male genitalia. A Merodon bequaerti, epandrium, lateral view B Merodon bequaerti, epandrium, 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 I Merodon sophron, hypandrium, 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, l–lingula, pl–posterior surstyle lobe. Scale bar: 0.2 mm.
(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.
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 I Merodon sophron, outer side, lateral view, female. Scale bar: 1 mm.
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 fascia of black pile between wing basis; scutum with two or more microtrichose vittae, anteriorly connected and posteriorly reaching the scutellum; 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 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 metafemur, slightly longer than pile on dorsal margin (Fig.
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.
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.
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.
Merodon bequaerti is distributed in north-western Africa (Algeria, Libya, Morocco, and Tunisia) (Fig.
Preferred environment: unimproved montane grassland, including open, grassy areas in pine forest or Mediterranean scrub. Flowers visited: no data. Flight period: February-June.
Holotype [original designation by
Algeria • 1 ♂; Kabylie, Tikjda; 36°27'00"N, 4°07'60"E; 28 Jun. 1954;
Libya • 1 ♀; 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.;
Morocco • 1 ♂; Moyen Atlas, Azrou; 33°25'48"N, 5°12'36"W; 15 Jun. 1928; R. Benoist leg.;
Tunisia • 1 ♂; Jundubah, 40 km W from Jendouba; 36°31'54"N, 8°28'25"E; 17 May 1988;
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.
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 disjunctus sp. nov., inner side, lateral view, male I 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.
Male genitalia. A Merodon defectus sp. nov., epandrium, lateral view B Merodon defectus sp. nov., 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 I 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, l–lingula, pl–posterior surstyle lobe. Scale bar: 0.2 mm.
Male. Head. Antennae black to dark brown; basoflagellomere elongated, 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.
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 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 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 incrassate, 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.
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, yellow, 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.
Female. Similar to the male except for normal sexual dimorphism and for the following characteristics: basoflagellomere ca. two times longer than wide (Fig.
Latin adjective defectus (reduced in size, smaller) refers to small basolateral protrusion (lateral hump) on posterior surstyle lobe.
Merodon defectus sp. nov. has been identified in western Turkey (Fig.
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 [
Holotype. Turkey • ♂; Bozdağ mountain, Near Bozdağ; 38°22'28"N, 28°04'38"E; 1140 m a.s.l.; 7 Jun. 2014; A. Vujić, J. Ačanski leg.;
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.
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 dorsomedial and dorsolateral including apex of basoflagellomere (Fig.
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.
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.
Male genitalia. A Merodon disjunctus sp. nov., epandrium, lateral view B Merodon disjunctus sp. nov., 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 kawamurae, hypandrium, lateral view. Abbreviations: al–anterior surstyle lobe, bp–basolateral protrusion, l–lingula, pl–posterior surstyle lobe, s–lateral sclerite of aedeagus. Scale bar: 0.2 mm.
Female. Similar to the male except for normal sexual dimorphism and for the following characteristics: basoflagellomere 1.6–1.8 times longer than wide, fossette large, dorsolateral (Fig.
The name derives from the Latin adjective disjunctus meaning separated, disconnected which pertains to the dichoptic eyes in the males.
Merodon disjunctus sp. nov. has so far only been recorded in Kyrgyzstan and Kazakhstan (Fig.
Preferred environment: no data. Flowers visited: no data. Flight period: May-July.
Holotype. Kyrgyzstan • ♂; 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.;
Kyrgyzstan • 1 ♂; 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.;
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 basoflagellomere (Fig.
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, female 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 I Merodon medium sp. nov., outer side, lateral view, male J Merodon medium sp. nov., outer side, lateral view, female. Scale bar: 1 mm.
(based on lectotype and additional specimens from the type locality, 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 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.
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 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 covered 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 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.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, ca. two times longer than wide, covered with short pile (Fig.
Female. Similar to the male except for normal sexual dimorphism and for the following characteristics: antennae with rounded tip, basoflagellomere 1.8–2 times longer than wide, fossette dorsolateral (Fig.
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, lateral view, female. Scale bar: 2 mm.
Merodon hirsutus is distributed in Israel, Syria, and south-eastern Turkey (Fig.
Preferred environment: no data. Flowers visited: no data. Flight period: March-June.
Described by
Israel • 1 ♀; Hefa, “Ma`yan Zevi” [Ma Yan Zevi]; 32°34'00"N, 34°55'60"E; 17 Apr. 1980;
Syria • 1 ♂; Jebel al Aqra; 35°55'18"N, 35°57'51"E; Jun. 1985; D. F. Leuthner leg.;
Turkey • 1 ♂; İçel, İcel-Taşucu, Silifke; 36°22'17"N, 33°54'54"E; 300 m a.s.l.; 17 Mar. 1984;
Lampetia micromegas
Hervé-Basin, 1929: 111 – syn. published by
Medium sized (7.7–11.2 mm), with olive-brown reflection; antennae reddish 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.
(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.
Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect yellow pile; scutum usually 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, dense pale yellow pile and grayish microtrichia; wings entirely 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.
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 specimen on 3); terga 2–4 each with a pair of white microtrichose, wide, usually oblique fasciae; 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.
Body parts, dorsal view. A Merodon trianguloculus sp. nov., thorax, male B Merodon trianguloculus sp. nov., thorax, female C Merodon trianguloculus sp. nov., abdomen, male D Merodon trianguloculus sp. nov., abdomen, female E Merodon kawamurae, abdomen, male F Merodon kawamurae, abdomen, female. Scale bar: 2 mm.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, covered with dense, short pile (Fig.
Female. Similar to the male except for normal sexual dimorphism and for the following characteristics: antennae with rounded tip, fossette dorsolateral (Fig.
Merodon kawamurae is known from Japan and China (Fig.
Preferred environment: no data. Flowers visited: no data. Flight period: April-May.
Merodon kawamurae was described after an unknown number of specimens from Kumamoto, Kyushu, Japan, leg. Kawamura. Matsumura’s type material 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
Paralectotypes (Lampetia micromegas). CHINA • 1 ♂; Chemo; 33°44'32"N, 103°23'45"E; 25 Apr. 1918;
CHINA • 1 ♀; Ningpo; 29°44'29"N, 121°06'02"E; 29 Apr. 1925; J. T. Chu leg.;
Large species (10.3–13 mm) with wide dark brown abdomen and yellow-orange maculae on lateral sides of tergum 2 (Fig.
Male. Head. Antennae black to dark brown; basoflagellomere elongated ca. 2.2 times as long as wide, and ca. 2.5 times as long as pedicel, concave dorsally 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.
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, anteriorly 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 covered 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.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, 1.5 times longer than wide, covered with dense, short pile (Fig.
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, l–lingula; arrow marks the hook-like extension in A. Scale bar: 0.2 mm.
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.
Medium (middle, center) refers to the species’ distribution, being the only taxon of the group found on Crete, in the middle of Mediterranean Sea.
Merodon medium sp. nov. is endemic to the Greek island of Crete (Fig.
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.
Holotype. Greece • ♂; Crete, Chania, Omalos plain; 35°19'21"N, 23°55'50"E; 28 May 2014; A. Vujić leg.;
Medium sized (10.2–10.8 mm), black and shiny species, covered with mostly black pile on scutum, terga and legs in both sexes (Fig.
Antenna. A Merodon nigropunctum sp. nov., outer side lateral view, male B Merodon nigropunctum 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.
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 dorsolateral; arista black and thickened at basal one third, covered with dense microtrichia, ca. two times as long as basoflagellomere (Fig.
Thorax. Scutum and scutellum black, shiny, covered with dense, erect, black pile; scutum without microtrichose vittae (Fig.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, ca. two times longer than wide, covered with dense, short pile (Fig.
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.
The name nigrocapillatus is derived from Latin adjective niger meaning black, dark and Latin noun capillatus meaning long-haired, referring to the long, black body pile of this species.
Merodon nigrocapillatus sp. nov. has only been recorded in Tajikistan (Fig.
Preferred environment: open areas at high altitudes, unimproved grassland (Fig.
Holotype. Tajikistan • ♂; Varzob, Kalon; 39°03'36"N, 68°52'12"E; 2440 m a.s.l.; 1–4 Jul. 2017; A. Barkalov leg.;
Medium sized (10.3 mm), bluish species (Fig.
Male. Head. Antennae black to dark brown; basoflagellomere narrow 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.
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.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, 1.5 times longer than wide, covered with dense, short pile (Fig.
Female. Unknown.
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.
Merodon nigropunctum sp. nov. was recorded at only one locality in Uzbekistan (Fig.
Preferred environment: no data. Flowers visited: no data. Flight period: May.
Holotype. Uzbekistan • ♂; Kadamžai, S of Fergana; 40°20'00"N, 71°47'39"E; 21 May 1980; Z. Padr leg.;
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 basoflagellomere (Fig.
Male. Head. Antennae black to dark brown; basoflagellomere elongated 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.
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 microtrichose vittae, anteriorly 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 covered with microtrichia; wing veins brown; calypteres yellowish; 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.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid in shape, 1.5 times longer than wide, covered with dense, short pile (Fig.
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.
(Fig.
Light micrographs of Merodon opacus sp. nov. puparium. A puparium in dorsal view B cephalopharyngeal 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 μm (B).
Male genitalia. A Merodon trianguloculus sp. nov., epandrium, lateral view B Merodon trianguloculus sp. nov., epandrium, ventral view C Merodon trianguloculus sp. nov., hypandrium, lateral view. Abbreviations: al–anterior surstyle lobe, bp–basolateral protrusion, l–lingula. Scale bar: 0.2 mm.
Latin adjective opacus (opaque, not transparent), pertains to the dark tergum 2, without reddish yellow lateral maculae.
Merodon opacus sp. nov. has been recorded on the Greek island of Lesvos and in western Turkey (Fig.
Preferred environment: forest/open ground; thermophilous and evergreen Quercus forest; Castanea forest, dry Pinus forest; unimproved grassland and tracksides (Fig.
Different types of habitats of Merodon serrulatus species group. A Lesvos (Greece), habitat of Merodon opacus sp. nov., Photograph by Ante Vujić B Siberia, Teletskoye Lake (Russia), habitat of Merodon serrulatus, Photograph by Jeroen van Steenis C Morocco, habitat of Merodon sophron, Photograph by Ante Vujić D Tajikistan, habitat of Merodon nigrocapillatus sp. nov., Photograph by Anatolij Barkalov.
Holotype. Greece • ♂; Lesvos; Polichnitos; 39°05'02"N, 26°09'13"E; 30 Apr. 2008; A. Vujić leg.;
Turkey • 3 ♀♀; 12 km SW of Muğla; 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 ♂♂; Muğla, University Campus; 37°09'42"N, 28°22'13"E; 700 m a.s.l.; 17–22 May 2011; M. Bartak, Kubik leg.; M. B. coll. 17927, 17928, 17930, 17931, 17937, 17938, 17940 to 17943, 17946, 17951, 17955 to 17960, 17965 to 17972 • 20 ♀♀; Muğla, University Campus; 37°09'42"N, 28°22'13"E; 700 m a.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 ♀; Muğla, 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 ♀♀; Muğla, 13 km NE pine wood; 37°14'50"N, 28°30'00"E; 1200 m a.s.l.; 23–27 Jun. 2015; M. Bartak, Kubik leg.; M. B. coll. 17924 to 17926 • 4 ♀♀; Bozdağ mountain, Near Bozdağ; 38°22'28"N, 28°04'38"E; 1140 m a.s.l.; 7 Jun. 2014; A. Vujić, J. Ačanski leg.;
Greece, Lesvos • 1 ♀; Agiassos; 39°03'00"N, 26°22'60"E; 28 May 2010; Horsfield leg.;
Large (9.5–11.6 mm) dark brown species with lack of microtrichose fasciae on terga 2–4 in males (Fig.
(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 microtrichia, 1.6 times as long as basoflagellomere (Fig.
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 microtrichose vittae, anteriorly 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 covered with microtrichia; wing veins brown; calypteres and halteres 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.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid in shape, 1.5 times longer than wide, covered with dense, short pile (Fig.
Female. Unknown.
Merodon sacki is known only from Spain (Fig.
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.
Holotype (original designation): male, “Holotypus Lampetia / sacki Paramononv, 1936 / G.V. POPOV des. 2007” [red label], “Lampetia / sacki n. sp. / ♂ Typus / Paramonov d.” [pink label handwritten], “Merodon / mir únbekannt” [yellow label handwritten], “14 VII 81” “Chiclana” [handwritten on the back side] (
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
Spain • 3 ♂♂; La Corte; 37°57'41"N, 6°49'09"W; 28 Apr. 2015; A. Vujić, D. Obreht leg.;
Merodon alexeji
Paramonov, 1925: 155 – syn. published in
Merodon lusitanicus
Hurkmans, 1993: 181 – syn. published in
Merodon tener Sack, 1913: 443 syn. nov.
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 yellow, 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.
(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.
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.
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 variability) (Fig.
Male genitalia. Apical part of anterior surstyle lobe triangular shape, 1.1–1.4 times longer than wide, covered with dense, short pile (Fig.
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.
There is some intra- and interpopulation variability in the morphological characters of Merodon serrulatus, which are summarized in Table
Character | Variability (intra- and/or interpopulation) | Intra | Inter |
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 in male | dorsal to lateral or dorsal and medial (Fig. |
+ | + Iberian populations with medial 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 scutum | from 2–4, posterior half dull without microtrichia | + | - |
black pile on scutum | few, or fascia of black pile between wing basis, or many black pile on scutum | + | - |
color of pile on metafemur in male | all yellowish to whitish or with many black in apical one third | + | + some Iberian populations with only pale pile |
length of pile on metafemur in male | one third to one fifth of width of metafemur | + | + in eastern populations (southern Russia to Siberia) longer, from one third to one fourth of width of metafemur |
color of knees, apex of tibiae and tarsi | from black to brown | + | - |
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 frons in female | from narrow unconnected to broad and connected near ocellar triangle | + | - |
color of pile on frons in female | almost all black to mostly whitish | + | - |
male genitalia | the shape of surstyle and size of area covered with dense short marginal setulae on anterior surstyle lobe (Figs |
+ | + in eastern populations (southern Russia to Siberia) basolateral protrusion less distinct (Fig. |
As shown in Fig.
Preferred environment: forest/open ground; thermophilous Quercus forest; Castanea forest (
Holotype of Merodon serrulatus [original designation in
Merodon alexeji: Described by
Merodon lusitanicus: Holotype [original designation by
Merodon tener: Described by
Croatia • 1 ♀; Velebit, Brušane; 44°29'55"N, 15°16'43"E; 600 m a.s.l.; 13 Jun. 1969;
France • 1 ♀; Languedoc-Roussillon, Corbieres, Carcassonne; 43°13'00"N, 2°21'00"E; 18 Jun. 1974;
Greece • 2 ♀♀; Mountain Taygetos; 22 km SW Sparta; 36°58'60"N, 22°24'09"E; 6 May 1990;
Italy • 1 ♀; 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.;
MONTENEGRO • 1 ♂; Lovćen, Lovćen 1; 42°22'59"N, 18°53'54"E; 17 May 2018; A. Vujić, A. Šebić, M. Ranković leg.;
Kazakhstan • 1 ♀; East Kazakhstan, Markakol’ District, 20 km N settlement Alekseevka, Souther slop of Matobaj Mountain range; 48°42'22"N, 85°57'00"E; 2318 m a.s.l.; 6 Jul. 1996; V. Zinchenko leg.;
NORTH MACEDONIA • 1 ♂; Kožuf, Golema poljana; 41°10'54"N, 22°12'05"E; 15 Jun. 1955;
Russia • 1 ♂; Sarepta, “Russia, RUS, Sarepta, now a suburb of Volgograd city (Christoph)”; 48°31'40"N, 44°29'01"E;
Spain • 1 ♂; Sierra Nevada, second valley; 37°06'10"N, 3°27'19"W; 1430 m a.s.l.; 17 Jun. 2014; A. Vujić, S. Radenković, C. Pérez-Bañón leg.;
Turkey • 5 ♂♂; “Kop Dağı geçidi” [Kop mountain pass], Bayburt; 40°15'00"N, 40°15'00"E; 16 Jul. 1992;
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 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.
(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.
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 fascia of short black pile and few black pile between wing basis; scutum with two microtrichose vittae, anteriorly connected and posteriorly reaching the scutellum; anterior half of 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, 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.
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.
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.
Merodon sophron is distributed in north-western Africa (Morocco) (Fig.
Preferred environment: forest/open ground; open areas in evergreen oak maquis, dry Pinus forest; unimproved grassland and tracksides (Fig.
Holotype [original designation by
Morocco • 1 ♂; Azrou; 30°40'00"N, 7°30'00"W; 31 May 1953; G. L. Spoek leg.;
Medium sized (7.5–11.6 mm), dark brown species with characteristic large silver microtrichose fasciae on terga 2–4 in males (Fig.
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.
Thorax. Scutum and scutellum black with bronze luster, covered with dense, erect, yellow pile; scutum with conspicuous silver microtrichose ornamentation (Fig.
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, approximately as long as wide, covered with dense, short pile (Fig.
Female. Similar to the male except for normal sexual dimorphism and for the following characteristics: basoflagellomere ca. 1.8 times longer than wide, fossette dorsolateral (Fig.
The name trianguloculus derives from the Latin adjective triangulus (triangular) and Latin noun loculus (spot) and describes the distinctive triangular silver pollinose fasciae on the abdomen.
Merodon trianguloculus sp. nov. was recorded only in Turkmenistan (Fig.
Preferred environment: open areas extending to the forest zone; unimproved 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.
Holotype. Turkmenistan • ♂; 120 km SW Geok-Tepe town; 38°10'31"N, 57°58'01"E; 11 May 1988; A. Barkalov leg.;
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 located in the Hungarian National Museum in Budapest, but the Diptera collection was destroyed by a fire in 1956. The description of
1 | Posterior part of mid coxa without long pile (Merodon avidus-nigritarsis lineage) | 2 |
– | Posterior part of mid coxa with long pile |
other Merodon lineages |
2 | Taxa with characteristic basolateral protrusion (lateral hump) on posterior surstyle lobe (as on Figs |
3 |
– | Species without basolateral protrusion (lateral hump) on posterior surstyle lobe in males and with different combinations of characters in females |
other species groups belonging to the Merodon avidus-nigritarsis lineage |
3 | Males | 4 |
– | Females | 19 |
4 | Eyes dichoptic (as on Figs |
5 |
– | Eyes holoptic (as on Fig. |
6 |
5 | Black species with predominantly black body pile, especially on thorax (Fig. |
Merodon nigrocapillatus sp. nov. |
– | Species with olive-brown reflection, predominantly covered with pale yellow pile; terga 2–4 with conspicuous lateral microtrichose fasciae (Fig. |
Merodon disjunctus sp. nov. |
6 | Bluish species (Fig. |
Merodon nigropunctum sp. nov. |
– | Dark brown species without dark macula on wings | 7 |
7 | Tergum 2 entirely dark brown to black (as on Fig. |
8 |
– | Tergum 2 with yellow-orange lateral maculae (at least small ones) (as on Fig. |
10 |
8 | Scutum without black pile, except few black setae at wing basis in some specimens; terga 2–4 with a conspicuous microtrichose fasciae (as on Fig. |
9 |
– | Scutum with black pile, at least on fascia between wing basis; terga 2–4 with a less conspicuous microtrichose fasciae (as on Fig. |
Merodon serrulatus (Wiedemann in Meigen, 1822) (part) |
9 | Dorsolateral pile on metafemur dense and longer (Fig. |
Merodon hirsutus Sack, 1913 |
– | Dorsolateral pile on metafemur shorter (Fig. |
Merodon opacus sp. nov. |
10 | Terga 3 and 4 with a pair of broad silver microtrichose maculae (Fig. |
Merodon trianguloculus sp. nov. |
– | Terga 3 and 4 without or with a less conspicuous pair of broad silver microtrichose fasciae (as on Fig. |
11 |
11 | Terga 3 and 4 without microtrichose fasciae | 12 |
– | Terga 3 and 4 with a pair of white microtrichose, oblique fasciae (as on Fig. |
13 |
12 | Metafemur strongly curved (Fig. |
Merodon sacki (Paramonov, 1936) |
– | Metafemur less curved (Fig. |
Merodon bequaerti Hurkmans, 1993 (part) |
13 | Antennae reddish yellow; basoflagellomere short and broad, ca. 1.2 times as long as wide, with large dorsal to dorsolateral fossette (Fig. |
Merodon kawamurae Matsumura, 1916 |
– | Antennae dark brown/black; basoflagellomere elongated; legs mostly black | 14 |
14 | Abdomen broad, tergum 2 at least 2.5 times wider than long (as on Fig. |
15 |
– | Abdomen narrower, tergum 2 ca. two times wider than long (as on Fig. |
16 |
15 | Pile on ventral margin of metafemur very short (Fig. |
Merodon medium sp. nov. |
– | Pile on ventral margin of metafemur long and dense (Fig. |
Merodon bequaerti Hurkmans, 1993 (part) |
16 | Male genitalia: basolateral protrusion (lateral hump) on posterior surstyle lobe reduced (Fig. |
Merodon defectus sp. nov. |
– | Male genitalia: basolateral protrusion (lateral hump) on posterior surstyle lobe well developed (as on Fig. |
17 |
17 | Medial fossette absent (as on Fig. |
18 |
– | Medial fossette present (Fig. |
Merodon serrulatus (Wiedemann in Meigen, 1822) |
18 | GenBank acc. no. MN623564-MN623581; distribution: Palaearctic, extending from France in the west to Turkey in the south-east, and to Siberia toward north-east (Fig. |
Merodon serrulatus (Wiedemann in Meigen, 1822) (part) |
– | GenBank acc. no. MN623540; distribution: north-west Africa (Fig. |
Merodon sophron Hurkmans, 1993 |
19 | Tergum 2 dark (as on Fig. |
20 |
– | Tergum 2 with reddish lateral maculae (as on Fig. |
22 |
20 | Black species (Fig. |
Merodon nigrocapillatus sp. nov. |
– | Species with olive-brown reflection, predominantly covered with pale pile | 21 |
21 | Pile on dorsolateral margin of metafemur long (Fig. |
Merodon hirsutus Sack, 1913 |
– | Pile on dorsolateral margin of metafemur short (Fig. |
Merodon opacus sp. nov. |
22 | Metafemur with long pile on the entire surface of ventral margin (as on 22B) (unknown female of Merodon sacki (Paramonov, 1936), probably keys out here) | 23 |
– | Metafemur with mostly short pile on ventral margin (as on Fig. |
27 |
23 | Scutum with characteristic silver microtrichose ornamentation (Fig. |
Merodon trianguloculus sp. nov. |
– | Scutum with indistinct microtrichose vittae; terga 3 and 4 with narrower lateral microtrichose fasciae | 24 |
24 | Scutum at wing basis with only yellowish pilosity; metafemur with long, dense dorsal pile (as on Fig. |
25 |
– | Scutum at wing basis with short black pile; metafemur with sparse dorsal pile (as on Fig. |
26 |
25 | Blackish species; terga 3 and 4 with broad lateral microtrichose fasciae; basoflagellomere dark brown to black; distribution: Kyrgyzstan and Kazakhstan (Fig. |
Merodon disjunctus sp. nov. |
– | Brownish species; terga 3 and 4 with narrower lateral microtrichose fasciae (Fig. |
Merodon kawamurae Matsumura, 1916 |
26 | Distribution: north-west Africa (Fig. |
Merodon bequaerti Hurkmans, 1993 |
– | Distribution: France | Merodon serrulatus (Wiedemann in Meigen, 1822) (part) |
27 | Females of these three species can be separated by distribution and genetic data: | |
– | GenBank acc. no. MN623540. Distribution: North-west Africa (Fig. |
Merodon sophron Hurkmans, 1993 |
– | GenBank acc. no. MN623564-MN623581. Distribution: Palaearctic, extending from Iberian Peninsula in the west to Turkey in the south-east and toward Siberia in the north-east (Fig. |
Merodon serrulatus (Wiedemann in Meigen, 1822) (part) |
– | Distribution: western Turkey (Fig. |
Merodon defectus sp. nov. |
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 characters (with gaps) were included in analyses for the D2−3 region of the 28S rRNA gene.
Both obtained phylogenetic trees (Maximum Parsimony, Fig.
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, Retention Index (RI) 65. Bootstrap support values are depicted near nodes (≥ 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.
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 differences 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 distinction between the species in each pair is based on characters with more subtle differences. These two pairs are M. bequaerti / M. sacki, with the metafemur incrassate and long pile on postero- and anteroventral surface of the metafemur, and M. hirsutus / M. opacus sp. nov., with tergum 2 dark, without yellow-orange lateral maculae in both sexes.
The remaining three species within the M. serrulatus species group are morphologically very similar to each other. Merodon defectus sp. nov. has subtle, but stable differences 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 (
In the present study we applied this integrative approach, i.e., to combine morphology, genetic data, and distribution, to support the taxonomic status and systematic decisions made for the M. 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. Discordance between morphological and molecular data has been observed in some previous studies concerning closely related taxa within the family Syrphidae (e.g.,
It is important to do further taxonomic research with the populations of Merodon serrulatus with high inter-population morphological and genetic variability. These populations 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 population structure that might contain evolutionary units at different levels of speciation.
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 (
The morphology of the puparium of M. 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 (
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 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: M. 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 snowdrop species], M. aureus species group and Crocus L. (Iridaceae) [
Being distributed from the Iberian Peninsula in the south-west, along the Mediterranean and Balkan Peninsula, through Turkey and southern Russia to Siberia and Mongolia 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, M. sacki has been found in southern Spain, M. 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. hirsutus found in south-eastern Turkey, Israel and Syria. The M. serrulatus species group includes two North-African species, i.e., M. 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 Japan. 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 (
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 particularly the case for the alpine areas, and especially in terms of the invertebrate fauna (
We thank the curators of the museums listed in the Materials and methods for facilitating 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 Technological Development of the Republic of Serbia, Grant Nos. OI173002 and III43002, the Provincial Secretariat for Science and Technological Development (0601-504/3), the H2020 Project “ANTARES” (664387) and the Scientific and Technological Research Council of Turkey (TÜBİTAK, project number: 213O243). The work of A.V. 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 Turkey by TÜBİTAK, and in Spain with permission N. Ref.: ENSN/FJSG/IMJ (232) (Junta de Andalucia, Consejería de Medio Ambiente y Ordenación del Territorio).
Roles of authors: AV, LL, SRad, CPB, AB, RH, GS and SRojo performed the sampling; 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, AŠ, 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.
Figure S1
Data type: type specimens' data
Explanation note: A Merodon bequaerti, holotype and labels B Merodon nigrocapillatus sp. nov., holotype and labels.
Figure S2
Data type: type specimens' data
Explanation note: A Merodon defectus sp. nov., holotype and labels B Merodon disjunctus sp. nov., holotype and labels.
Figure S3. Merodon medium sp. nov., holotype and labels
Data type: type specimens' data
Explanation note: Merodon medium sp. nov., holotype and labels.
Figure S4
Data type: type specimens' data
Explanation note: A Merodon nigropunctum sp. nov., holotype and labels B Merodon opacus sp. nov., holotype and labels
Figure S5. Merodon trianguloculus sp. nov., holotype and labels
Data type: type specimens' data
Figure S6
Data type: type specimens' data
Explanation note: A Merodon tener, lectotype and labels B Merodon sacki, holotype and labels.
Figure S7
Data type: phylogenetic data
Explanation note: Maximum likelihood tree based on analysis of combined COI mitochondrial 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).
Table S1. Data for the specimens used in the molecular analysis, including GenBank accession numbers
Data type: molecular specimens' dataset