Research Article |
Corresponding author: Andrijana Andrić ( andrijana.andric@biosense.rs ) Academic editor: Bradley Sinclair
© 2023 Ante Vujić, Snežana Radenković, Anatolij Barkalov, Nataša Kočiš Tubić, Laura Likov, Tamara Tot, Grigory Popov, Alex Prokhorov, Ebrahim Gilasian, Shehzad Anjum, Mihajla Djan, Banafsha Kakar, Andrijana Andrić.
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, Radenković S, Barkalov A, Kočiš Tubić N, Likov L, Tot T, Popov G, Prokhorov A, Gilasian E, Anjum S, Djan M, Kakar B, Andrić A (2023) Taxonomic revision of the Merodon tarsatus species group (Diptera, Syrphidae). Arthropod Systematics & Phylogeny 81: 201-256. https://doi.org/10.3897/asp.81.e93570
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The phytophagous hoverfly genus Merodon Meigen, 1803 (Diptera, Syrphidae) presently comprises 205 described species distributed in the Palaearctic and Afrotropical regions, and it is differentiated into multiple species groups. In this work, we revise the Merodon tarsatus species group, providing an illustrated key to species, a detailed discussion on relevant taxonomic characters, and a morphological diagnosis. We summarize morphological characteristics of the 12 species in the M. tarsatus group, together with available distributional data. Moreover, we provide descriptions for five new species, namely Merodon angustitarsis Vujić & Gilasian sp. nov., M. dumosus Vujić, Likov & Radenković sp. nov., M. latiantennatus Vujić, Popov & Prokhorov sp. nov., M. namaghijamii Vujić, Likov & Radenković sp. nov. and M. pakistanicus Vujić, Likov & Radenković sp. nov. Additionally, we describe one more species, Merodon nigroalatus Vujić, Likov & Radenković sp. nov., that belongs to the same lineage and is related to the Merodon tarsatus group based on molecular markers and structure of male genitalia, but lacks modifications of the metatarsus and sternum 4 that are the main diagnostic features of the group. Furthermore, we redefine and redescribe seven taxa of the M. tarsatus group. Following detailed analyses of type material, we revise the status of all available taxa related to M. tarsatus and we propose the following new synonyms: M. ankylogaster Hurkmans, 1993 syn. nov. (junior synonym of M. hypochrysos Hurkmans, 1993); M. persicus Hurkmans, 1993 syn. nov. (junior synonym of M. oidipous Hurkmans, 1993); and M. xanthipous Hurkmans, 1993 syn. nov. (junior synonym of M. marginicornis Hurkmans, 1993). We apply Maximum Parsimony and Maximum Likelihood analyses of the 5′ end of the mitochondrial COI gene sequences to genetically characterize species in the M. tarsatus group and the related species, M. nigroalatus Vujić, Likov & Radenković sp. nov.
COI gene, distributional data, hoverflies, morphology, new species, new synonyms
The phytophagous hoverfly genus Merodon Meigen, 1803 (Diptera: Syrphidae: Eristalinae: Merodontini) is distributed across the Palaearctic and Afrotropical regions and presently comprises 205 described species (
In summarizing previously published data (
The highest species diversity of Merodon has been recorded in the Mediterranean region (
The Merodon tarsatus species group is the only group of the genus with limited distribution in the Middle East and Central Asia that lacks European representatives. It was first established by
Here, we present a taxonomic review of the Merodon tarsatus species group based on a detailed examination of material gathered from our long-term research on the genus Merodon. Our aims are to: 1) review material of this group stored in several major entomological institutions and private collections; 2) define and describe the taxa within the M. tarsatus species group, including new species; 3) provide diagnostic characters of all species and complete a key for the M. tarsatus group; and 4) conduct DNA sequence analysis of the 5′ region of the mitochondrial COI gene for nine species of the M. tarsatus group. We additionally processed a species close to the M. tarsatus group that belongs to the same lineage.
In the present study specimens belonging to the Merodon tarsatus species group, as well as additional species close to it, Merodon nigroalatus Vujić, Likov & Radenković sp. nov. belonging to avidus-nigritarsis lineage, were examined. In total 1368 specimens (1365 belonging to the M. tarsatus group and three specimens of M. nigroalatus Vujić, Likov & Radenković sp. nov.) from Afghanistan, Iran, Israel, Kazakhstan, Kyrgyzstan, Pakistan, Tajikistan, Turkey, Turkmenistan, and Uzbekistan were studied. The material examined in this study are deposited in the following entomological collections:
Daniele Sommaggio collection, Bologna, Italy (D.S. coll.);
Dieter Doczkal collection, Munich, Germany (D.D. coll.);
Entomological Museum of Isparta, Isparta, Turkey (EMIT);
innish Museum of Natural History, University of Helsinki, Helsinki, Finland (
All type specimens of species in the Merodon tarsatus group were examined. Within the “Type material examined” sections below, we provide complete data for each studied specimen. For holotypes, information on general condition is described. The original label data has been given verbatim for the type specimens. The contents of each label are presented in quotation marks (“”), with a slash (/) denoting separate lines within a label. For non-type specimens, data on specimens is presented in the sections entitled “Additional material examined”.
Differential diagnoses are given for all studied species. We describe five new species in full and redescribe a further seven species based on terminology following
Photographs were composed with software CombineZ (
Information about distribution is given for all studied species. DIVA-GIS software version 7.5.0 (
Laboratory process. The genomic DNA of each specimen was extracted from two or three legs using a slightly modified SDS extraction protocol (
Data analysis. The obtained COI gene sequences were edited for base-calling errors using BioEdit 7.2.5. (
The Merodon avidus-nigritarsis lineage contains medium to large species (11–20 mm) usually with white pollinose vittae on scutum and white pollinose fasciae on terga; anterior anepisternum bare ventrally to the postpronotum; abdomen elongate, usually narrow and tapering, longer than scutum and scutellum together; posterior part of mesocoxa usually without long pile (with exception of M. eumerusi); basoflagellomere usually at most twice as long as wide; legs without additional calcar,spina(e) (with exception of M. eumerusi) or tubercle; male genitalia (as in Figs
Male genitalia, hypandrium, lateral view. A, B M. tarsatus, C M. turkestanicus, D M. namaghijamii Vujić, Likov & Radenković sp. nov. (A, C, D) hypandrium, (B) aedeagus. Abbreviations: ea – ejaculatory apodeme, l – lingula, s – lateral sclerite of aedeagus, x – maximal width of ejaculatory apodeme, y – basal width of hypandrium. Scale bar: 0.5 mm.
The Merodon tarsatus species group belongs to the avidus-nigritarsis lineage on the basis of the absence of long pile posteriorly on the mesocoxa. This group contains small to large species (6–13 mm) characterized in males with sternum 4 medially clearly divided with membranous structure and with lateral tubercles or laminate extensions on posterior margin (Fig.
The Merodon tarsatus group consists of seven described and five undescribed species. This group of species is geographically restricted to the Middle East and Central Asia (Figs
M. aff. tarsatus
1 (
IRAN: Māzandarān Province, Baladeh District, Nesen; 36°14’24”N, 51°27’27”E; 2920 m a.s.l.
Holotype
: ♂; HMIM [specimen dry pinned, in very good condition with genitalia in separate microvial]. Original labels: “Mazandaran-Baladeh / Nesan / N 36 14 24.2 E 051 27 27.0 / 2920m. / Gilasian / 9.VI.2006” [left part of the label vertical], “02532”. — Paratypes: IRAN • 1 ♀; Yazd Province, Taft County, Sānij; 31°33’36”N, 54°01’46”E; 3020 m a.s.l.; 15 May 2007; Gilasian E. leg.; HMIM 04461 • 2 ♂♂; N. Iran, Alborz (mountain range), Tochal; 35°53’50”N, 51°25’05”E; 2000–2500 m a.s.l.; 29–30 Jun. 1973; Loc. no. 261 Exp. Nat. Mus. Praha; HMIM 02530, 02536 • 1 ♀; same data as for preceding; HMIM 02537 • 1 ♂; N. Iran, Alborz (mountain range), Tochal; 35°53’50”N, 51°25’05”E; 2000–2500 m a.s.l.; 29–30 Jun. 1973; Loc. no. 261 Exp. Nat. Mus. Praha;
Metabasotarsomere. A–F M. smirnovi, G M. turkestanicus, H–K M. oidipous, L–N M. tarsatus. (A, B, H, I, L, M) male, (C–G, J, K, N) female; (A, E, F, H, K, N) lateral view, (B–D, I, J, L) ventral view, (G, M) dorsal view. Arrows mark length of brush-like area of dense pile. Scale bars: 0.5 mm (A–L, N), 1 mm (M).
IRAN • 1 ♀; Fārs Province, Road Chiraz (Shiraz)-Kazeroun, Fort Sine-Sefid; 29°30’N, 52°E; 29 Apr. 1937; Brandt F.H. leg.; Paratype of Merodon ankylogaster;
Medium sized (8–10 mm), medium long pilose, dark species, with olive-brown reflection (Figs
Metabasotarsomere. A–D M. dumosus Vujić, Likov & Radenković sp. nov., E M. latiantennatus Vujić, Popov & Prokhorov sp. nov., F–H M. marginicornis. (A, B, E–G) male, (C, D, H) female; (A, G) dorso-lateral view, (B, D, F) ventral view, (C, E, H) lateral view. Arrow marks length of brush-like area of dense pile. Scale bar: 0.5 mm.
Male clearly differs from Merodon rufitarsis by a longer basotarsomere of metaleg in M. angustitarsis Vujić & Gilasian sp. nov. (Fig.
Metabasotarsomere of male. A–C M. namaghijamii Vujić, Likov & Radenković sp. nov., D M. pakistanicus Vujić, Likov & Radenković sp. nov., E–F M. turkestanicus. (A) dorsal view, (B, D, E) lateral view, (C, F) ventral view. Arrows mark length of brush-like area of dense pile. Scale bars: 0.5 mm (A–D, F), 0.75 mm (E).
MALE. Head: Antenna reddish-brown; basoflagellomere (Fig.
The name “angustitarsis”, as arbitrary combination, derives from the Latin adjective “angustus” meaning narrow, and the noun “tarsus” referring to the fifth part (terminal segments) of the leg. This describes the distinctive narrow shape of the basotarsomere of the metaleg.
Merodon angustitarsis Vujić & Gilasian sp. nov. occurs in Iran and in southern Turkmenistan (Fig.
M. aff. tarsatus 2 (
ISRAEL: Mount Hermon [Hebrew: Har Hermon]; 2000 m a.s.l.
Holotype
: ♂; TAUI 04907 [specimen dry pinned, in very good condition with genitalia in separate microvial]. Original labels: “ISRAEL: Har / Hermon, 2000m / 12.vi.1996 / A. Friedberg”, “04907”. — Paratypes: IRAN • 1 ♀; Kermān Province, Bāft County, Deh Sard; 28°45’22”N, 56°31’22”E; 2280 m a.s.l.; 22 Apr. 2006; Gilasian E. leg.; HMIM 02571 • 1 ♂; Kermān Province, Bāft County, Ghale Asgar; 29°30’13”N, 56°38’19”E; 2740 m a.s.l.; 4 May 2007; Gilasian E., Nematian M. leg.; HMIM 02570 • 1 ♀; same data as for preceding; HMIM 02534 • 1 ♂; Kermān Province, Rayen District, Kuh-e Hazaran; 29°30’42”N, 57°16’18”E; 3800 m a.s.l.; 25 May 1978; Warncke K. leg.;
Medium to large (10–13 mm), long pilose, dark species, with olive-brown reflection (Figs
Male clearly differs from similar species, Merodon latiantennatus Vujić, Popov & Prokhorov sp. nov. and M. marginicornis by ventrolateral row of strong setae on basotarsomere of all legs (Fig.
Male. Head: Antenna reddish to dark brown; basoflagellomere (Fig.
The name “dumosus” is a Latin adjective in masculine, derived from the noun “dumus” meaning “overgrown with thorns”. This term refers to a row of strong setae on the basotarsomere.
Merodon dumosus Vujić, Likov & Radenković sp. nov. occurs in Iran and Israel (Fig.
Hurkmans & Friedberg identified this species in an unpublished manuscript.
Merodon hypochrysos Hurkmans, 1993: 165. Type locality: Turkey, Adiyaman province (holotype).
Merodon ankylogaster Hurkmans, 1993: 169. Type locality: Iran, Fārs province. Syn. nov.
Turkey, Adiyaman province; Iran, Fārs province.
Merodon ankylogaster Hurkmans: Original description was based on male holotype and female paratype from the same locality (
IRAN • ♂; Fārs Province, Road Chiraz (Shiraz)-Kazeroun, Fort Sine-Sefid; 29°35’N, 52°E; 29 Apr. 1937; Brandt F.H. leg.;
Merodon hypochrysos Hurkmans: Original description was based on holotype and 20 paratypes from Hakkari province in Turkey (
Hurkmans designated Merodon ankylogaster as a member of the “tarsatus group”. The male holotype of M. ankylogaster, designated by
IRAN • 1 ♂; Kermān Province, Bāft County, Ghale Asgar; 29°30’13”N, 56°38’19”E; 2740 m a.s.l.; 3 May 2007; Gilasian E., Nematian M. leg.; HMIM • 5 ♂♂, 2 ♀♀; Fārs Province, 15 km S of Dasht-e Arjan; 29°33’09”N, 51°56’22”E; 2261 m a.s.l.; 2–6 May 2016; Obořil M. leg.; J.H. coll. • 4 ♂♂, 2 ♀♀; Fārs Province, Dasht-e Arjan; 29°37’48”N, 51°54’43”E; 2040 m a.s.l.; 5 May 2016; Kafka M. leg.; M. B. coll. • 2 ♂♂, 2 ♀♀; Kohgiluyeh & Boyer-Ahmad Province, Yāsuj, Sarab-e Taveh; 30°29’24”N, 51°39’29”E; 2390 m a.s.l.; 4 May 2016; Kafka M. leg.; M. B. coll. • 1 ♀; Isfahān Province (Esfahān Province), Semirom County, Kommeh; 31°01’01”N, 51°35’28”E; 2760 m a.s.l.; 12 May 2007; Gilasian E. leg.; HMIM • 5 ♂♂; Lorestān Province, Dorud County, Lanjabad; 33°25’08”N, 48°59’10”E; 950 m a.s.l.; 10 May 2016; Kafka M. leg.; M. B. coll. • 1 ♂; Lorestān Province, Dorud County, Lanjabad; 33°25’59”N, 49°01’44”E; 670 m a.s.l. (+/-300 m); 11 May 2016; Kafka M. leg.; M. B. coll. • 2 ♀♀; Lorestān Province, Dorud County, Lanjabad; 33°25’59”N, 49°01’44”E; 1670 m a.s.l.; 10–11 May 2016; Kafka M. leg.; M. B. coll. • 2 ♂♂; Tehran Province, Damāvand Village, Damāvand Mt area, 75 km NE Teheran (Tehran); 35°58’32”N, 52°06’20”E; 27 Jul. 1976; Lavalle A.G. leg.; USNM ENT 00036574, USNM ENT 00036575, 05120 (
Small sized (6–8 mm), short to medium long pilose, dark species, with olive-brown reflection (Fig.
Differs from other species from the Merodon tarsatus group by shorter pilosity on metafemur (especially dorsally), with pile shorter than base of metatibia in lateral view (Figs
Male. Head: Antenna dark brown; basoflagellomere (Fig.
Merodon hypochrysos occurs in the region of the Middle East, including Iran, Israel and southeastern Turkey (Fig.
M. aff. tarsatus
3 (
AFGHANISTAN: Kabul Province, Paghman District, Paghman; 2500–2600 m a.s.l.
Holotype : ♂; SIZK 25352 [specimen dry pinned in good condition, the right wing is glued on locality label, right meso- and metaleg are missing, taken for molecular analysis, genitalia in separate microvial]. Original labels: “AFGHANISTAN / KABUL PROV., PAGHMAN / H=2500–2600m / 9.06.2016 O. PAK”, “TS797”, “25352”. — Paratypes: AFGHANISTAN • 1 ♀; Kabul Province, Paghman District, Paghman; 2600 m a.s.l.; 2 Jun. 2010; SIZK 25353 • 2 ♀♀; same data as for preceding; 2700 m a.s.l.; SIZK • 1 ♀; same data as for preceding; Pak O. leg. • 1 ♀; same data as for preceding; 1 Jul. 2013 • 1 ♀; same data as for preceding; 4 Jul. 2013 • 1 ♀; same data as for preceding; 2 Jul. 2013; Skrylnik Yu. leg. • 3 ♀♀; same data as for preceding; 2500–2600 m a.s.l.; 9 Jun. 2016; Pak O. leg.
A–I Metaleg of male, lateral view J, K metatarsus, dorsal view. A, J M. latiantennatus Vujić, Popov & Prokhorov sp. nov., B M. oidipous, C M. rufitarsis, D M. smirnovi, E–G M. tarsatus, H, I M. turkestanicus, K M. dumosus Vujić, Likov & Radenković sp. nov. Abbreviation: mt – metatrochanter. Scale bars: 1 mm (A–I), 0.5 mm (J, K).
Small (7–8 mm), dark species, with bluish reflection (Fig.
Similar to Merodon marginicornis from which male differs in shorter and less spiky basoflagellomere (Fig.
Male. Head: Antenna orange; basoflagellomere (Fig.
The name “latiantennatus”, as arbitrary combination, is derived from the Latin adjectives “latus” meaning “wide, broad” and “antennatus” meaning “having antenna”, referring to the shape of the male basoflagellomere.
Merodon latiantennatus Vujić, Popov & Prokhorov sp. nov. has so far only been recorded in eastern Afghanistan (Fig.
Merodon marginicornis Hurkmans, 1993: 166. Type locality: Iran, Fārs province, Shirāz (holotype).
Merodon xanthipous Hurkmans, 1993: 175. Type locality: Iran, Fārs province, Shirāz. Syn. nov.
Iran, Fārs province, Shirāz.
Merodon xanthipous Hurkmans: Original description was based on male holotype and female paratype from the localities close to the locality of holotype of Merodon Merodon (
Merodon marginicornis Hurkmans: Original description was based on one male specimen designated as holotype (
Hurkmans designated M. xanthipous as a member of “tarsatus group”. Both taxa belong to the same species and based on prior citation of M. marginicornis in the same publication, M. xanthipous becomes a junior synonym.
Additional material examined. IRAN • 1 ♂; Fārs Province, Dasht-e Arjan; 29°33’07”N, 51°56’31”E; 2260 m a.s.l. (+/–300 m); 4 May 2016; Kafka M. leg.; M. B. coll. • 4 ♂♂, 1 ♀; same data as for preceding; 5 May 2016 • 3 ♂♂; Fārs Province, 15 km S of Dasht-e Arjan; 29°33’09’’N, 51°56’22”E; 2261 m a.s.l.; 2–6 May 2016; Obořil M. leg.; J.H. coll. 18257, 18258, 18259 • 2 ♀♀; same data as for preceding; J.H. coll. 18269, 18275 • 1 ♂; Fārs Province, Sepidān; 30°17’13”N, 51°58’15”E; 2540 m a.s.l., 8 May 2007; Gilasian E., Nematian M. leg.; HMIM 04460 • 1 ♀; Kermān Province, Māhān District, Bolbolouyeh Village; 30°09’37”N, 57°22’10’’E; 2430 m a.s.l.; 29 Apr. 2007; Gilasian E., Nematian, M. leg.; HMIM 04459 • 1 ♂, 1 ♀; Kohgiluyeh & Boyer-Ahmad Province, Yāsuj, Sarab-e Taveh; 30°29’24”N, 51°39’29”E; 2390 m a.s.l.; 4 May 2016; Kafka M. leg.; M. B. coll. • 1 ♀; Isfahān Province (Esfahān Province), Semirom County, Kommeh; 31°01’01”N, 51°35’28”E; 2760 m a.s.l.; 12 May 2007; Gilasian E. leg.; HMIM 02543 • 1 ♀; Chāhārmahāl & Bakhtiāri Province, 30 km SE of Lordegān; 31°21’00”N, 51°09’00”E; 1900 m a.s.l.; 31 May 2014; Halada J. leg.; M. B. coll. • 1 ♂; Alborz Province, Karaj County, 10 km N of Gachsar; 36°09’00”N, 51°18’00”E; 2300–2700 m a.s.l.; 7 Jun. 2014; Halada J. leg.; M. B. coll. 10433. — PAKISTAN • 1 ♀; Balochistan Province, Ziarat District, 35 km W of Ziarat; 30°23’14”N, 67°20’22”E; 11 May 1984; McGinley R.J. leg.; 05117, USNM ENT00036567 (
Small to medium sized (7–11 mm), medium long pilose, dark species, with olive-brown reflection (Figs
Male similar to Merodon latiantennatus Vujić, Popov & Prokhorov sp. nov. from which differs in more elongated basoflagellomere with pointed apex (Fig.
Male. Head: Antenna reddish-brown; basoflagellomere (Fig.
Merodon marginicornis was recorded in Iran, southern Turkmenistan, and western Pakistan (Fig.
Hurkmans treated many species groups at the same time in the monograph (
M. aff. tarsatus
4 (
IRAN: Razavi Khorasan Province, Torbat-e Jām County, Bezd village, 20 km SW of Torbat-e-Jām; 35°21’00”N, 60°44’00”E; 1274 m a.s.l.
Holotype
: ♂;
Male genitalia. A–C M. angustitarsis Vujić & Gilasian sp. nov., D–F M. rufitarsis, G–I M. hypochrysos. (A, B, D, E, G, H) epandrium, (C, F, I) hypandrium. (A, C, D, F, G, I) lateral view, (B, E, H) ventral view. Abbreviations: al – anterior surstyle lobe, c – cercus, ea – ejaculatory apodeme, l – lingula, pl – posterior surstyle lobe. Scale bar: 0.5 mm.
Medium sized (10 mm), long pilose, dark species, with olive-brown reflection (Fig.
Similar to Merodon turkestanicus from which differs in shape and size of basotarsomere of metaleg, broader in M. namaghijamii Vujić, Likov & Radenković sp. nov. (Fig.
Description. Male. Head: Antenna dark brown to black; basoflagellomere (Fig.
The new species “namaghijamii” was named after the famous Persian writer, poet: Ahmad Ibn Abolhasan Jāmi-e Nāmaghi-e Torshizi. He was born in Namagh, Iran (1048) and died in Torbat-e Jām, Iran (1141). The holotype was collected from the county named Torbat-e-Jām, located near the border of Iran and Afghanistan. This name was proposed by Hussein Sadeghi Namaghi from Iran. A noun in the genitive case.
Merodon namaghijamii Vujić, Likov & Radenković sp. nov. has only been recorded in northeastern Iran so far (Fig.
Merodon oidipous Hurkmans, 1993: 171. Type locality: Turkey, Hakkari province (holotype).
Merodon persicus Hurkmans, 1993: 171. Type locality: Iran, Fārs province. Syn. nov.
Turkey, Hakkari province; Iran, Fārs province.
Merodon persicus Hurkmans: Original description was based on one male specimen designated as holotype (
Male genitalia. A–C M. dumosus Vujić, Likov & Radenković sp. nov., D–F M. latiantennatus Vujić, Popov & Prokhorov sp. nov., G–I M. marginicornis. (A, B, D, E, G, H) epandrium, (C, F, I) hypandrium; (A, C, D, F, G, I) lateral view, (B, E, H) ventral view. Abbreviations: al – anterior surstyle lobe, c – cercus, ea – ejaculatory apodeme, l – lingula, pl – posterior surstyle lobe. Scale bar: 0.5 mm.
Hurkmans designated Merodon persicus as a member of “tarsatus group”. Specimens described as M. oidipous and M. persicus belong to one species and based on prior citation of M. oidipous in the same publication, M. persicus becomes a junior synonym.
IRAN • 2 ♂♂, 2 ♀♀; Fārs Province, Dasht-e Arjan; 29°33’07”N, 51°56’31”E; 2260 m a.s.l.; 5 May 2016; Kafka M. leg.; M. B. coll. • 1 ♂; Fārs Province, 15 km S of Dasht-e Arjan; 29°33’09”N, 51°56’22”E; 2261 m a.s.l.; 2–6 May 2016; Obořil M. leg.; J.H. coll. • 3 ♀♀; same data as for preceding; J.H. coll. 18265, 18273, 18275 • 1 ♂; Fārs Province, 10 km E Kazeroun; 29°34’00”N, 51°52’00”E; 1300 m a.s.l.; 23 May 2014; Halada J. leg.; M. B. coll. • 1 ♀; Fārs Province, 1 km W of Sangar; 29°59’50”N, 52°08’07”E; 2093 m a.s.l.; 4–5 May 2016; Obořil M. leg.; J.H. coll. 18268 • 4 ♂♂, 3 ♀♀; Kohgiluyeh & Boyer-Ahmad Province, Yāsuj, Sarab-e Taveh; 30°29’24”N, 51°39’29”E; 2390 m a.s.l.; 4 May 2016; Kafka M. leg.; M. B. coll. • 1 ♂; Kohgiluyeh & Boyer-Ahmad Province, 20 km S of Yāsuj (Jásúdž); 30°29’26”N, 51°39’27”E; 2144 m a.s.l.; 30 May 2015; Baňař P. leg.; J.H. coll. 18266 • 1 ♀; same data as for preceding; J.H. coll. 18272 • 1 ♂; Isfahān Province (Esfahān Province), Semirom County, Kommeh; 31°01’01”N, 51°35’28”E; 2760 m a.s.l.; 12 May 2007; Gilasian E. leg.; HMIM 04462.
Medium sized (8–11 mm), long pilose, dark species with olive-brown to bluish reflection (Fig.
Similar to Merodon smirnovi and M. tarsatus from which differs in smaller ventral brush-like area of dense pile on basotarsomere of metaleg, limited to basal 1/3 (Fig.
Male. Head: Antenna reddish-brown to dark brown; basoflagellomere (Fig.
Merodon oidipous occurs in southern Iran and southeastern Turkey (Fig.
M. aff. tarsatus
5 (
PAKISTAN: Balochistan Province, Qilla Saifullah District, Kan Mehtarzai.
Holotype : ♂; NARC 18120 [specimen dry pinned, the right wing, the head, antero-dorsal part of mesonotum and mesoleg without tarsomeres are glued on locality label, metaleg taken for molecular analysis, genitalia in separate microvial]. Original labels: “Kan Mwhterzai / (Qilla Saifulah) / 27-v-2017 / Banafsha”, “10 [serial number] ♂”, “18120”, “TS512”.
Medium sized (10 mm), long pilose, dark species, with olive-brown reflection; antennae black, basoflagellomere 1.8 times as long as wide (Fig.
Differs from Merodon namaghijamii Vujić, Likov & Radenković sp. nov. and M. turkestanicus by holoptic eyes, eye contiguity about 10 facets long (Fig.
Male. Head: Antenna black; basoflagellomere (Fig.
The new species “pakistanicus” was named after the country of origin of the holotype (Pakistan).
Merodon pakistanicus Vujić, Likov & Radenković sp. nov. has only been recorded in western Pakistan so far (Fig.
Merodon fulcratus
Tajikistan, the Alajsski Mountains.
Holotype
: TAJIKISTAN • ♂; original label: ‘Alai mont’
KAZAKHSTAN • 1 ♀; Aksu-Zhabagly; 42°17’33”N, 70°40’15”E; 1400–1500 m a.s.l.; 4 Jun. 1996; Dolin W. leg.; M. H. coll. 02575. — KYRGYZSTAN • 1 ♀; 40 km S of Jany Bazar; 41°42’00”N, 71°06’00”E; 8–11 Jun. 1995; Halada J. leg.; AM-05-216;
Medium sized (8–11 mm), medium long pilose, dark species, with olive-brown reflection (Figs
Similar to Merodon angustitarsis Vujić & Gilasian sp. nov. from which differs by absence of ventrolateral row of setae on basotarsomere of metaleg, present in M. angustitarsis Vujić & Gilasian sp. nov. (Fig.
Male. Head: Antenna reddish-yellow to brown; basoflagellomere (Fig.
Merodon rufitarsis occurs in northwestern Kyrgyzstan, Tajikistan, eastern Turkmenistan and southern Kazakhstan (Fig.
This species was described as subspecies of Merodon fulcratus, but
Merodon smirnovi Paramonov, 1927: 76 (= 320).
Turkestan, Tashkent.
Lectotype (designated here in order to fix identity of the species): UZBEKISTAN • 1 ♂; Turkestan, d. Tashkent, Ak-Tash; 21 Jun. 1925; Smirnov E.S. leg.; SIZK [specimen dry pinned in good condition, left fourth and fifth tarsomeres of metaleg missing, genitalia in separate microvial]. Original labels: “Ak-Tash / d. Tashkent / Turkestan / 21.VI.25. Smirnov l.” [yellowish, pale violet ink, black-bordered label], “Merodon / "smirnovi n. sp. / ♂. Cotypus / Paramonov. det” [pink double black-bordered, pale violet ink label], “Lectotypus / Merodon smirnovi Param., 1927 G.V. Popov des. 2007” [red label], “02568”. — Paralectotype (designated here): UZBEKISTAN • 1 ♀; Turkestan, d. Tashkent, Ak-Tash; 21 Jun. 1925; Smirnov E.S. leg.; SIZK [specimen dry pinned in very good condition]. Original label: “Ak-Tash / d. Tashkent / Turkestan / 21.VI.25. Smirnov l.” [yellowish, pale violet ink, black-bordered label], “ Merodon / "smirnovi n. sp. / ♀ Cotypus / Paramonov. det” [pink double black-bordered, pale violet ink label], “Paralectotypus / Merodon Merodon smirnovi Param., 1927 G.V. Popov des. 2007” [red label], “02569”.
KAZAKHSTAN • 1 ♀; Fabritchny, 40 km E of Alma Ata; 43°13’21”N, 77°20’54”E; 23 Jun. 1992; Halada M. leg.;
Medium sized to large (9–13 mm), long pilose, dark species with olive-brown reflection (Fig.
Similar to Merodon tarsatus from which differs in sternum 2 with medial patch of long white pile (Fig.
Male. Head: Antenna dark brown to reddish-brown; basoflagellomere (Fig.
Merodon smirnovi occurs in Kyrgyzstan, southeastern Kazakhstan, northeastern Uzbekistan (near borders with Kyrgyzstan and Kazakhstan) and western Tajikistan (Fig.
Original description was based on two syntypes from Uzbekistan (
Merodon tarsatus Sack, 1913: 437.
Tajikistan, Pamir.
Lectotype
: syntype cited as “holotype” in
AFGHANISTAN • 1 ♀; Badakschan, Schiva high steppe; 7 Jul. 1953; Klapperich J. leg.; Bańkowska R. det. as Merodon Merodon Paramonov;
Medium sized to large (9–13 mm), long pilose, dark species, with olive-brown reflection (Figs
Similar to Merodon smirnovi from which differs in sternum 2 with pile uniformly distributed (Fig.
Male. Head: Antenna black to dark brown (Fig.
Merodon tarsatus occurs in Tajikistan, Kyrgyzstan and northeastern Afghanistan (Fig.
Original description was based on two syntypes from Tajikistan (
Merodon turkestanicus Paramonov, 1927: 77 (= 321).
Uzbekistan, Tashkent.
Holotype : UZBEKISTAN • ♂. Original labels: “N 339”, “d. Tashkent / Turkestan / 12.V.15.” [yellowish black-bordered, pale violet ink], “Merodon / turkestanicus / n. sp. ♂ Typus / Paramonov. det” [pink double black-bordered, violet ink], “Holotypus / Merodon Merodon Par., 1927 G.V.Popov des. 2007” [red].
KAZAKHSTAN • 1 ♂; Krasnogorski, Dzhushy-Dala, near Anrakhay; 43°41’53”N, 74°49’33”E; 28 Apr. 1956; Marikovsky P. leg.;
Medium sized (9–11 mm), long pilose, dark species, with olive-brown reflection (Fig.
Differs from males of Merodon smirnovi, M. tarsatus and M. pakistanicus Vujić, Likov & Radenković sp. nov. by dichoptic eyes or shortly connected by distance of 1–5 facets long in M. turkestanicus (Fig.
Male. Head: Antenna dark brown; basoflagellomere (Fig.
Maximum-Likelihood 5′COI tree of the Merodon tarsatus species group based on the Tamura-Nei model. Tree with highest log likelihood (–2540.4813) is shown. A discrete Gamma distribution was used to model evolutionary rate differences among sites (5 categories (+G, parameter = 0.1619)). Bootstrap values ≥ 50 are presented near nodes.
Merodon turkestanicus occurs in Tajikistan, Southeastern Kazakhstan, northeastern Uzbekistan and southern Turkmenistan (Kopet-Dag mountain range) (Fig.
Original description was based on a holotype from Uzbekistan (
1. |
Basotarsomere of metaleg ventrally with well-defined, brush-like area of dense pile (as in Figs |
2 |
– | Basotarsomere of metaleg ventrally without well-defined, brush-like area of dense pile (as in Fig. |
11 |
2. |
Basoflagellomere stubby (ca. 1.7 times as long as wide), triangular, orange (Fig. |
Merodon latiantennatus Vujić, Popov & Prokhorov sp. nov. |
– | Basoflagellomere more elongated (at least 2 times as long as wide), differently shaped (as in Fig. |
3 |
3. | Sternum 2 with medial patch of long whitish pile (Fig. |
Merodon smirnovi Paramonov, 1927 |
– | Sternum 2 with uniformly distributed long whitish pile (as in Fig. |
4 |
4. |
Brush-like area of setae on basotarsomere of metaleg limited to basal 1/3 ventrally (Fig. |
Merodon oidipous Hurkmans, 1993 |
– | Brush-like area of setae on basotarsomere of metaleg extended at least 1/2 ventrally (as in Figs |
5 |
5. | Male genitalia with enlarged, oval anterior surstyle lobe, longer than posterior surstyle lobe (Fig. |
Merodon dumosus Vujić, Likov & Radenković sp. nov. |
– | Male genitalia with anterior surstyle lobe as long as or shorter than posterior surstyle lobe (as in Fig. |
6 |
6. | Basotarsomere of metaleg elongated, less expanded, less than 2 times as broad as second tarsomere in lateral view (Fig. |
Merodon marginicornis Hurkmans, 1993 |
– | Basotarsomere of metaleg expanded, more than 2 times as broad as second tarsomere in lateral view, without spine-like setae along ventrolateral margin (as in Figs |
7 |
7. |
Eyes holoptic, eye contiguity more than 6 facets long (Figs |
8 |
– | Eyes dichoptic (Fig. |
10 |
8. | Basotarsomere of metaleg strongly expanded, about 3 times broader than second tarsomere (Fig. |
Merodon tarsatus Sack, 1913 |
– | Basotarsomere of metaleg less expanded, about 2 times broader than second tarsomere (as in Figs |
9 |
9. | Metafemur swollen, 3 times broader than metatibia in ventral view; male genitalia: ejaculatory apodeme longer than broad (Fig. |
Merodon pakistanicus Vujić, Likov & Radenković sp. nov. |
– | Metafemur not swollen, 2 times broader than metatibia in ventral view; male genitalia: ejaculatory apodeme almost as long as wide (Fig. |
Merodon turkestanicus Paramonov, 1927 (in part) |
10. | Basotarsomere of metaleg less expanded, about 2 times broader than second tarsomere (Fig. |
Merodon turkestanicus Paramonov, 1927 (in part) |
– | Basotarsomere of metaleg more expanded, about 2.5 times broader than second tarsomere (Fig. |
Merodon namaghijamii Vujić, Likov & Radenković sp. nov. |
11. |
Metafemur and metatibia broader, covered with dense pilosity (Fig. |
Merodon rufitarsis Sack, 1913 |
– | Metafemur and metatibia narrower, covered with less dense pilosity (Fig. |
12 |
12. | Basotarsomere of metaleg elongated, 3.5 times longer than wide (Fig. |
Merodon angustitarsis Vujić & Gilasian sp. nov. |
– | Basotarsomere of metaleg short and flat, 1.5 times longer than wide dorsally with parallel margins (Fig. |
Merodon hypochrysos Hurkmans, 1993 |
(Females of Merodon pakistanicus Vujić, Likov & Radenković sp. nov. and M. namaghijamii Vujić, Likov & Radenković sp. nov. are unknown.)
1. | Metafemur with short pilosity, shorter than base of metatibia in lateral view, except a few long pile dorsally (Fig. |
Merodon hypochrysos Hurkmans, 1993 |
– | Metafemur with longer pilosity, particularly dorsally, longer than base of metatibia in dorsal view (as in Figs |
2 |
2. | Metafemur and metatibia broad, with long and dense yellow pilosity (Fig. |
Merodon rufitarsis Sack, 1913 |
– | Species with different combination of characters | 3 |
3. |
Basotarsomere of metaleg narrow and elongated, 3.5 times or more longer than wide (as in Fig. |
4 |
– | Basotarsomere of metaleg broader and shorter, less than 3.3 times longer than wide (as in Fig. |
8 |
4. | Second tarsomere of metaleg longer, about half length of basotarsomere (Fig. |
Merodon angustitarsis Vujić & Gilasian sp. nov. |
– | Second tarsomere of metaleg shorter, less than half length of basotarsomere (as in Fig. |
5 |
5. | Larger species (10–13 mm); longer spine-like setae within ventrolateral row of setae very strong and distinct (Fig. |
Merodon dumosus Vujić, Likov & Radenković sp. nov. |
– | Smaller species (7–11 mm); longer spine-like setae within ventrolateral row of setae less distinct (as in Figs |
6 |
6. | Longer spine-like setae within ventrolateral row of setae indistinct or absent (as in Fig. |
7 |
– | Longer spine-like setae within ventrolateral row of setae distinct (Fig. |
Merodon marginicornis Hurkmans, 1993 (in part) |
7. | Basoflagellomere brown to black; distribution: Turkmenistan, Tajikistan, Kazakhstan, Uzbekistan | Merodon turkestanicus Paramonov, 1927 |
– | Basoflagellomere orange-brown (Figs |
Merodon latiantennatus Vujić, Popov & Prokhorov sp nov. |
8. | Sternum 2 medially with pile at least two times longer than laterally (Fig. |
Merodon smirnovi Paramonov, 1927 |
– | Sternum 2 with pilosity uniform in length (as in Fig. |
9 |
9. | Some longer spine-like setae within ventrolateral row of setae very distinct (Fig. |
Merodon marginicornis Hurkmans, 1993 (in part) |
– | All spine-like setae within ventrolateral row of setae equal in length (as in Fig. |
10 |
10. | Sternum 2 clearly shorter than sternum 3; basotarsomere of metaleg from ventral view usually medially narrowed and twisted (Fig. |
Merodon oidipous Hurkmans, 1993 |
– | Sternum 2 as long as or slightly shorter than sternum 3; basotarsomere of metaleg from ventral view usually medially not narrowed and twisted; basoflagellomere brown to dark brown (Fig. |
Merodon tarsatus Sack, 1913 |
(Females of M. namaghijamii Vujić, Likov & Radenković sp. nov., and M. pakistanicus Vujić, Likov & Radenković sp. nov. probably also keyed here.)
Some species of the Merodon tarsatus group, such as M. dumosus Vujić, Likov & Radenković sp. nov., M. marginicornis, M. oidipous, M. smirnovi and M. tarsatus, show variability in range of their body size, with 4–5 mm differences between minimum and maximum sized specimens. Most of the species have variable color of basoflagellomere, from dark brown to reddish-brown, and in some species, such as M. oidipous and M. marginicornis, the basotarsomere of the metaleg varies in color, from black to reddish. The basotarsomere of the metaleg in M. smirnovi in both sexes (Fig.
Country distribution of species of the Merodon tarsatus group (bold font = type locality).
species | Afghanistan | Iran | Israel | Kazakhstan | Kyrgyzstan | Pakistan | Tajikistan | Turkey | Turkmenistan | Uzbekistan |
hypochrysos | X | X | X | |||||||
angustitarsis sp. nov. | X | X | ||||||||
dumosus sp. nov. | X | X | ||||||||
namaghijamii sp. nov. | X | |||||||||
latiantennatus sp. nov. | X | |||||||||
marginicornis | X | X | X | |||||||
pakistanicus sp. nov. | X | |||||||||
oidipous | X | X | ||||||||
rufitarsis | X | X | X | X | ||||||
smirnovi | X | X | X | X | ||||||
tarsatus | X | X | X | |||||||
turkestanicus | X | X | X | X |
Most specimens of Merodon turkestanicus possess dichoptic eyes (Fig.
M. aff. tarsatus
6 (
PAKISTAN: Khyber Paktunkhawa Province, Haripur District, Tehsil, Khanur Nikrian.
Holotype : ♂; NARC 05922 [specimen dry pinned, in good condition, both wings are damaged, genitalia in separate microvial]. Original labels: “Nikrian, Khan Pur / 29.iii.[20]13 / A. Haq”, “6B”, “05922”. — Paratypes: PAKISTAN • 1 ♂; Khyber Paktunkhawa Province, Haripur District, Tehsil, Khanur Nikrian; 29 Mar. 2013; Haq A. leg.; NARC 05923 • 1 ♀; same data as for preceding; NARC 05933.
Species with unique combination of morphological characters in Merodon avidus-nigritarsis lineage: abdomen with red markings (Figs
Merodon nigroalatus Vujić, Likov & Radenković sp. nov., male. A body, dorsal view; B head, frontal view; C wing, dorsal view; D basoflagellomere, lateral view; E protibia and protarsus, dorsal view; F sterna, ventral view; G metafemur and metatibia, lateral view; H metatibia and metatarsus, dorso-lateral view; I metatarsus, lateral view. Scale bars: 1 mm (A–C, E–H), 0.25 mm (D), 0.5 mm (I).
Male. Head: Antenna reddish-yellow; basoflagellomere (Fig.
The name “nigroalatus”, as arbitrary combination, is derived from Latin adjectives “niger” meaning black and “alatus” meaning winged. This describes the distinctive dark wing of the species.
Merodon nigroalatus Vujić, Likov & Radenković sp. nov., male genitalia. A epandrium, lateral view; B posterior surstyle lobe, dorsal view; C hypandrium, lateral view. Abbreviations: al – anterior surstyle lobe, c – cercus, d – membrane connecting two lobes of posterior surstyle lobe, l – lingula, pl – posterior surstyle lobe. Two lobes of posterior surstyle lobe marked with arrows. Scale bar: 0.5 mm.
Merodon nigroalatus Vujić, Likov & Radenković sp. nov. has only been recorded in northern Pakistan so far (Fig.
Mimics bees of the genus Sphecodes Latreille (Hymenoptera: Halictidae). This species has a unique position inside the genus based on very distinct morphological characters. Based on molecular marker and structure of the male genitalia it is related to the Merodon tarsatus group, but because of absence of the main diagnostic characters of the group, such as structure of sternum 4 and basotarsomere of metaleg, it should be kept as unplaced species inside the avidus-nigritarsis lineage.
The genetic analyses involved 38 sequences of 5′COI gene including outgroups, with a total of 621 nucleotide positions in the final dataset, of which 113 were parsimony informative. Maximum-Parsimony (MP) analysis resulted in 16 equally parsimonious trees of 351 step lengths, consistency index (Ci) = 63 and retention index (Ri) = 75 (Fig.
Three species within the Merodon tarsatus group display very distinct diagnostic features: M. hypochrysos has a short basoflagellomere with a concave dorsal margin and a rounded apex, as well as a flattened male basotarsomere of the metatarsus with parallel margins as observed from the dorsal view; M. rufitarsis is characterized by all tarsi being bright yellow; and M. smirnovi possesses unique pilosity of sternum 2, with a medial patch of long white pile, and an elongated basoflagellomere that is more than 2.5 times as long as wide. Notably, despite well documented variability in body size, basoflagellomere color and tarsi color within this species group, width of the metaleg basotarsomere relative to the second tarsomere proved a reliable character for species separation in both sexes of M. dumosus Vujić, Likov & Radenković sp. nov., M. marginicornis, M. oidipous, M. smirnovi and M. tarsatus.
We have characterized three species based on a strongly expanded basotarsomere of the metaleg, which was ≥ 3 times broader than the second tarsomere in male M. oidipous, M. smirnovi and M. tarsatus. In contrast, the basotarsomere of the male metaleg in M. namaghijamii Vujić, Likov & Radenković sp. nov., M. pakistanicus Vujić, Likov & Radenković sp. nov. and M. turkestanicus is narrower, approximately twice as broad as the second tarsomere. A third group of species possesses a narrow basotarsomere of the male metaleg, being only ~ 1.5 times (or less) as broad as the second tarsomere, namely M. angustitarsis Vujić & Gilasian sp. nov., M. dumosus Vujić, Likov & Radenković.sp. nov., M. latiantennatus Vujić, Popov & Prokhorov sp. nov. and M. marginicornis.
In addition to our assessment of morphological characters, we conducted genetic analyses based on sequences of the 5′ end of the mitochondrial COI gene. Many previous studies have highlighted the need for concurrent analyses of different character sets to assess diversity and relationships within Merodon (e.g.,
Our genetic analysis clustered M. smirnovi, M. pakistanicus Vujić, Likov & Radenković sp. nov., M. namaghijamii Vujić, Likov & Radenković sp. nov. and M. tarsatus together and these were clearly distinct genetically from the other species we analyzed. This outcome is congruent with their distinct morphological characters, i.e., broad basotarsomere of the metaleg and similar male genitalia.
Merodon tarsatus displays notable intraspecific morphological diversity. The male of this species presents variability in the shape of the metafemur, from strongly curved to almost straight. We obtained DNA sequences for specimens possessing either one of these morphological states, but all specimens resolved as a single clade on our 5′ COI gene trees (Figs
Merodon marginicornis and two other clearly morphologically distinct species (M. oidipous and M. latiantennatus Vujić, Popov & Prokhorov sp. nov.) formed a single genetic clade, with no clear molecular separation between the specimens of M. marginicornis and M. latiantennatus Vujić, Popov & Prokhorov sp. nov.. Similarly, it has been previously noted that COI gene sequences failed to distinguish morphologically well-defined species (
Our genetic data and the structure of male genitalia revealed a close relationship between M. nigroalatus Vujić, Likov & Radenković sp. nov. and the M. tarsatus species group. Nevertheless, given its lack of the primary morphological diagnostic characters of the M. tarsatus group—sternum 4 without lateral tubercles or laminate extensions on the posterior margin in males, and the metabasotarsomere not being expanded in both sexes—we retain M. nigroalatus Vujić, Likov & Radenković sp. nov. as a separate taxon within the avidus-nigritarsis lineage.
The genus Merodon is distributed throughout the Palaearctic and Afrotropical regions. Notably, the avidus-nigritarsis lineage is limited to the Palaearctic, but the M. tarsatus species group within that lineage lacks European representatives. The Mediterranean region displays the highest species diversity of Merodon, with the Anatolian peninsula considered a center of diversity and endemism (
Indeed, four of the twelve species of this group considered herein have only been recorded within the Middle East (Figs
The overall distribution of the Merodon tarsatus species group encompasses two biodiversity hotspots (sensu
Both of these biodiversity hotspots lie within the Irano-Turanian region, one of the largest floristic regions in the world, which exhibits a heterogeneous topography shaped by its complex tectonic history, thus fostering diverse isolated habitats that give rise to high species diversity and endemism (
Nevertheless, the Irano-Turanian region remains understudied, particularly its alpine areas and invertebrate fauna (
The immature stages of the avidus-nigritarsis lineage have only been described for two species to date: M. avidus (M. avidus species group) described from larvae and puparia in the bulbs of Ornithogalum L. (Asparagaceae) and surrounding soil (
Overall, the results of the present study reaffirm previous conclusions emphasizing the importance of underexplored regions, such as the Irano-Anatolian and Mountains of Central Asia biodiversity hotspots, in hosting high Merodon diversity (
We thank the curators of the museums listed in the Material and Methods for facilitating visits and loans for the study of specimens in their care. The authors owe a special debt of gratitude to Professor Hussein Sadeghi Namaghi from Iran for sending a specimen, suggesting the name of new species Merodon namaghijamii and for all his help relating to the data of collecting this specimen. We sincerely thank Volodymyr L. Perepetchayenko, Oleg V. Pak (Donetsk, Ukraine), and Yury E. Skrylnik (Kharkiv, Ukraine) for Afghan material loaned for the study. The photographs of the M. latiantennatus sp. nov. type locality were given by Yury E. Skrylnik. We thank John O’Brien for English proofreading, and the reviewers whose constructive input helped improve this manuscript. The authors acknowledge financial support of the Ministry of Education, Science and Technological Development of the Republic of Serbia (Grant No. 451-03-9/2021-14/200125, Grant No. 451-03-9/2021-14/200358, Grant No. 451-03-68/2022-14/200125 and Grant No. 451-03-68/2022-14/200358). The authors declare that no competing interests exist.
Table S1
Data type: .xlsx
Explanation note: List of molecularly analyzed samples with GenBank accession numbers (in boldface: newly generated sequences within this study).