Corresponding author: Laura Likov (
Academic editor: X. Mengual
The phytophagous hoverfly genus
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
The phytophagous hoverfly genus
The taxonomic status and identification of many
Most recent publications pertaining to the genus
The
In this study, we present a taxonomic review of the
Most of the recently collected specimens were sampled by sweep net. Further specimens of the
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.
The type material of all described species of the
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,
Sampling
A targeted search for immature
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
Basoflagellomere 1.5–2.2 times as long as wide, usually obviously concave dorsally (as in Fig.
The
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.
Distribution map of
Metaleg, lateral view.
Male genitalia.
Abdomen of males.
(based on the type material and additional specimens).
Antenna.
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.
Preferred environment: unimproved montane grassland, including open, grassy areas in pine forest or Mediterranean scrub. Flowers visited: no data. Flight period: February-June.
Algeria • 1 ♂; Kabylie, Tikjda;
Libya • 1 ♀; Tripolitania, Garian;
Morocco • 1 ♂; Moyen Atlas, Azrou;
Tunisia • 1 ♂; Jundubah, 40 km W from Jendouba;
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.
Metaleg, lateral view.
Male genitalia.
Abdomen, dorsal view.
Head of male.
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
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.
Latin adjective
Preferred environment: forest/open ground; thermophilous and evergreen
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.
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.
The name derives from the Latin adjective
Preferred environment: no data. Flowers visited: no data. Flight period: May-July.
Kyrgyzstan • 1 ♂; Tchatkal Valley, 4 SW Ajgyr-Dzhal vill.;
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.
(based on lectotype and additional specimens from the type locality, Syria, and Israel).
Head, dorsal view.
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.
Abdomen.
Preferred environment: no data. Flowers visited: no data. Flight period: March-June.
Described by
Israel • 1 ♀; Hefa, “Ma`yan Zevi” [Ma Yan Zevi];
Syria • 1 ♂; Jebel al Aqra;
Turkey • 1 ♂; İçel, İcel-Taşucu, Silifke;
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
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.
Male genitalia. Apical part of anterior surstyle lobe rhomboid shape, covered with dense, short pile (Fig.
Preferred environment: no data. Flowers visited: no data. Flight period: April-May.
CHINA • 1 ♀; Ningpo;
Large species (10.3–13 mm) with wide dark brown abdomen and yellow-orange maculae on lateral sides of tergum 2 (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.
Medium (middle, center) refers to the species’ distribution, being the only taxon of the group found on Crete, in the middle of Mediterranean Sea.
Preferred environment: forest/open ground; evergreen oak forest, dry
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.
Head of male, dorsal view.
Antenna.
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.
The name
Preferred environment: open areas at high altitudes, unimproved grassland (Fig.
Medium sized (10.3 mm), bluish species (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.
Male genitalia.
The word
Preferred environment: no data. Flowers visited: no data. Flight period: May.
Holotype. Uzbekistan • ♂; Kadamžai, S of Fergana;
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.
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.
(Fig.
Light micrographs of
SEM micrographs of
Male genitalia.
Latin adjective
Preferred environment: forest/open ground; thermophilous and evergreen
Different types of habitats of
Turkey • 3 ♀♀; 12 km SW of Muğla;
Greece, Lesvos • 1 ♀; Agiassos;
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).
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.
Preferred environment: forest/open ground; open areas in evergreen oak forest (
Holotype (original designation): male, “Holotypus Lampetia / sacki Paramononv, 1936 / G.V. POPOV des. 2007” [red label], “
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;
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).
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.
There is some intra- and interpopulation variability in the morphological characters of
Inter- and intrapopulation variability of
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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
Holotype of
Croatia • 1 ♀; Velebit, Brušane;
France • 1 ♀; Languedoc-Roussillon, Corbieres, Carcassonne;
Greece • 2 ♀♀; Mountain Taygetos; 22 km SW Sparta;
Italy • 1 ♀; Sicily, Etna, Rifugio Filiciusa;
MONTENEGRO • 1 ♂; Lovćen, Lovćen 1;
Kazakhstan • 1 ♀; East Kazakhstan, Markakol’ District, 20 km N settlement Alekseevka, Souther slop of Matobaj Mountain range;
NORTH MACEDONIA • 1 ♂; Kožuf, Golema poljana;
Russia • 1 ♂; Sarepta, “Russia, RUS, Sarepta, now a suburb of Volgograd city (Christoph)”;
Spain • 1 ♂; Sierra Nevada, second valley;
Turkey • 5 ♂♂; “Kop Dağı geçidi” [Kop mountain pass], Bayburt;
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).
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
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.
Preferred environment: forest/open ground; open areas in evergreen oak maquis, dry
Holotype [original designation by
Morocco • 1 ♂; Azrou;
Medium sized (7.5–11.6 mm), dark brown species with characteristic large silver microtrichose fasciae on terga 2–4 in males (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.
The name
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.
The identity of
1 | Posterior part of mid coxa without long pile ( |
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– | Posterior part of mid coxa with long pile |
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2 | Taxa with characteristic basolateral protrusion (lateral hump) on posterior surstyle lobe (as on Figs |
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– | Species without basolateral protrusion (lateral hump) on posterior surstyle lobe in males and with different combinations of characters in females |
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3 | Males |
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– | Females |
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4 | Eyes dichoptic (as on Figs |
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– | Eyes holoptic (as on Fig. |
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5 | Black species with predominantly black body pile, especially on thorax (Fig. |
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– | Species with olive-brown reflection, predominantly covered with pale yellow pile; terga 2–4 with conspicuous lateral microtrichose fasciae (Fig. |
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6 | Bluish species (Fig. |
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– | Dark brown species without dark macula on wings |
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7 | Tergum 2 entirely dark brown to black (as on Fig. |
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– | Tergum 2 with yellow-orange lateral maculae (at least small ones) (as on Fig. |
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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. |
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– | Scutum with black pile, at least on fascia between wing basis; terga 2–4 with a less conspicuous microtrichose fasciae (as on Fig. |
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9 | Dorsolateral pile on metafemur dense and longer (Fig. |
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– | Dorsolateral pile on metafemur shorter (Fig. |
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10 | Terga 3 and 4 with a pair of broad silver microtrichose maculae (Fig. |
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– | Terga 3 and 4 without or with a less conspicuous pair of broad silver microtrichose fasciae (as on Fig. |
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11 | Terga 3 and 4 without microtrichose fasciae |
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– | Terga 3 and 4 with a pair of white microtrichose, oblique fasciae (as on Fig. |
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12 | Metafemur strongly curved (Fig. |
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– | Metafemur less curved (Fig. |
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13 | Antennae reddish yellow; basoflagellomere short and broad, ca. 1.2 times as long as wide, with large dorsal to dorsolateral fossette (Fig. |
|
– | Antennae dark brown/black; basoflagellomere elongated; legs mostly black |
|
14 | Abdomen broad, tergum 2 at least 2.5 times wider than long (as on Fig. |
|
– | Abdomen narrower, tergum 2 ca. two times wider than long (as on Fig. |
|
15 | Pile on ventral margin of metafemur very short (Fig. |
|
– | Pile on ventral margin of metafemur long and dense (Fig. |
|
16 | Male genitalia: basolateral protrusion (lateral hump) on posterior surstyle lobe reduced (Fig. |
|
– | Male genitalia: basolateral protrusion (lateral hump) on posterior surstyle lobe well developed (as on Fig. |
|
17 | Medial fossette absent (as on Fig. |
|
– | Medial fossette present (Fig. |
|
18 | GenBank acc. no. |
|
– | GenBank acc. no. |
|
19 | Tergum 2 dark (as on Fig. |
|
– | Tergum 2 with reddish lateral maculae (as on Fig. |
|
20 | Black species (Fig. |
|
– | Species with olive-brown reflection, predominantly covered with pale pile |
|
21 | Pile on dorsolateral margin of metafemur long (Fig. |
|
– | Pile on dorsolateral margin of metafemur short (Fig. |
|
22 | Metafemur with long pile on the entire surface of ventral margin (as on 22B) (unknown female of |
|
– | Metafemur with mostly short pile on ventral margin (as on Fig. |
|
23 | Scutum with characteristic silver microtrichose ornamentation (Fig. |
|
– | Scutum with indistinct microtrichose vittae; terga 3 and 4 with narrower lateral microtrichose fasciae |
|
24 | Scutum at wing basis with only yellowish pilosity; metafemur with long, dense dorsal pile (as on Fig. |
|
– | Scutum at wing basis with short black pile; metafemur with sparse dorsal pile (as on Fig. |
|
25 | Blackish species; terga 3 and 4 with broad lateral microtrichose fasciae; basoflagellomere dark brown to black; distribution: Kyrgyzstan and Kazakhstan (Fig. |
|
– | Brownish species; terga 3 and 4 with narrower lateral microtrichose fasciae (Fig. |
|
26 | Distribution: north-west Africa (Fig. |
|
– | Distribution: France |
|
27 | Females of these three species can be separated by distribution and genetic data: | |
– | GenBank acc. no. |
|
– | GenBank acc. no. |
|
– | Distribution: western Turkey (Fig. |
|
The final aligned and pruned dataset including two-gene data matrix (
Both obtained phylogenetic trees (Maximum Parsimony, Fig.
Strict consensus tree of 41 most parsimonious trees from the analysis of combined
The
The remaining three species within the
Using different methodologies to assess various aspects of the diversity of the genus
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
It is important to do further taxonomic research with the populations of
One of the main reasons for the gap in extant knowledge on the immature stages of
The morphology of the puparium of
A single larva of
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,
The genus
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.
not treated here
not treated here
part: Iberian populations
unknown female of Merodon nigropunctum sp. nov. probably keys out here
Figure S1
type specimens' data
Figure S2
type specimens' data
Figure S3.
type specimens' data
Figure S4
type specimens' data
Figure S5.
type specimens' data
Figure S6
type specimens' data
Figure S7
phylogenetic data
Maximum likelihood tree based on analysis of combined
Table S1. Data for the specimens used in the molecular analysis, including GenBank accession numbers
molecular specimens' dataset