Corresponding author: Ante Vujić (
Academic editor: X. Mengual
The genus
Vujić A, Radenković S, Likov L, Veselić S (2021) Taxonomic complexity in the genus
The genus
The genus
In Europe,
The genus
Nowadays, with the advent of molecular and morphometric techniques, an integrative taxonomic framework has become the standard to study the taxonomy of genus
The aim of this work is to summarize the knowledge from recent taxonomic and systematic revisions, to help taxonomists to have a central reference for the recent published literature, and to present an identification key for the identification of lineages, species groups, species subgroups and unplaced species of
A total of 255 species belonging to the tribe
Morphological terminology follows
Male genitalia were extracted from dry specimens previously relaxed in a humidity chamber. After genitalia were pulled out with a hook-tipped entomological pin, they were cleared by boiling in warm 10% potassium hydroxide (KOH) for 3–5 min. Acetic acid was then used to neutralize the KOH during 5 s, and genitalia were immersed briefly in ethanol to remove the acid. Prepared genitalia were stored in microvials containing glycerol.
Photographs were taken using a Leica DFC 320 digital camera, attached to a Leica MZ16 stereomicroscope and Nikon Coolpix D7100 digital camera attached to a Nikon SMZ 745T stereomicroscope. Digital photographs were stacked using CombineZ software (
Based on
As mentioned earlier, there are five monophyletic lineages within the genus
In this section and sections below, we provide several identification keys to the 24 species groups, two species subgroups and 10 unplaced species within the genus
1 | Mesocoxa without long pile posteriorly (Suppl. material |
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– | Mesocoxa with at least a few long pile posteriorly (Suppl. material |
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2 | Mesocoxa with more than 10 long pile posteriorly |
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– | Mesocoxa with a few long pile posteriorly (usually five to seven, or less) (Suppl. material |
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3 | Basoflagellomere elongated, twice as long as wide, narrowed in apical third (Suppl. material |
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– | Basoflagellomere less than half as long as wide, narrowed in apical half (Suppl. material |
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4 | Anterior anepisternum with bare area ventral to postpronotum (Suppl. material |
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– | Anterior anepisternum with many long pile ventral to postpronotum (Suppl. material |
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5 | Postpronotum usually brown or yellow-reddish. Male genitalia: anterior surstyle lobe with curved distal prolongation (Suppl. material |
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– | Postpronotum black. Male genitalia: anterior surstyle lobe undeveloped (Suppl. material |
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6 | Lateral sclerite of the aedeagus gradually tapered, with the tip curved (Suppl. material |
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– | Lateral sclerite of the aedeagus hammer-like (Suppl. material |
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1 | Postpronotum, lateral sides of scutum and face yellowish (Suppl. material |
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– | Postpronotum, lateral sides of scutum and face black or dark |
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2 | Pro- and mesolegs strongly modified (Suppl. material |
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– | Pro- and mesolegs with usual shape |
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3 | Pilosity on the posterior part of abdomen (at least tergum 4) denser and strikingly golden to reddish-yellow (as in Suppl. material |
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– | Pilosity on the posterior part of abdomen not denser compared to the anterior part (Suppl. material |
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4 | Large (11–17 mm) bumble bee-like species with dense and long body pilosity (Suppl. material |
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– | Species with different characters |
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5 | Medium to large sized species (9–13 mm) with black, bronze lustre terga (lack pollinose fasciate maculae), except a pair of small, orange, lateral markings on tergum 2 (Suppl. material |
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– | Species with different characters; usually with pollinose fasciate maculae on terga |
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6 | Female (genitalia not visible) |
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– | Male (genitalia visible externally) |
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7 | Metaleg with some modifications on metatrochanter, metafemur and/or metatibia; male genitalia often with hook-like posterior surstyle lobe or cercus with prominence(s) |
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– | Metaleg usually without modifications (exception is |
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8 | Metatrochanter with blunt calcar apically covered with long pile (Suppl. material |
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– | Metatrochanter with sharp apical calcar (Suppl. material |
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9 | Metatibia narrow, not swollen apically (Suppl. material |
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– | Metatibia incrassate apically (Suppl. material |
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10 | Scutum usually with five distinct pollinose vittae (Suppl. material |
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– | Scutum with indistinct pollinose vittae; terga 2–4 without or with narrower pollinose fasciate maculae (Suppl. material |
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1 | Female (genitalia not visible) |
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– | Male (genitalia visible externally) |
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2 | Metatrochanter with calcar (Suppl. material |
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– | Metatrochanter rounded and smooth, without calcar (Suppl. material |
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3 | Hypandrium strongly modified, sinuous in apical half, with subapical ctenidium and stitched theca (cuticle looks like it is sewed) (Suppl. material |
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– | Hypandrium different, but with apical ctenidium and without stitch on theca (as in Suppl. material |
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4 | Pedicel elongated, approximately as long as basoflagellomere (relation pedicel basoflagellomere = 0.9 : 1.1) (Suppl. material |
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– | Pedicel shorter than basoflagellomere (as in Suppl. material |
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5 | Yellow-grey pilosity on terga more dense and striking laterally, as well as on pollinose fasciate maculae of terga 2 and 3, and on tergum 4 (Suppl. material |
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– | Terga evenly covered with upstanding, dense pilosity (can be differently coloured) (Suppl. material |
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6 | Pedicel shorter than basoflagellomere (Suppl. material |
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– | Pedicel elongated, approximately as long as basoflagellomere (relation pedicel : basoflagellomere = 0.9 : 1.1) (Suppl. material |
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7 | Apical triangular lamina on metafemur weakly serrated, usually with distinct apical dens (Suppl. material |
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– | Apical triangular lamina on metafemur markedly serrated (Suppl. material |
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8 | Terga without pollinose fasciate maculae, with dense puncta (Suppl. material |
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– | Terga 2–4 (at least 2) usually with pollinose medial fasciate maculae, with less dense puncta (Suppl. material |
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9 | Tergum 4 covered with short adpressed pile (Suppl. material |
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– | Tergum 4 covered with longer semi-erect pile, longer than basoflegellomere (Suppl. material |
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1 | Inner side of metafemur with a row of spinae (Suppl. material |
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– | Inner side of metafemur without a row of spinae |
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2 | Terga partly brown, reddish or yellow |
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– | Terga black |
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3 | Females (genitalia not visible) |
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– | Males (genitalia visible externally) |
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4 | Male genitalia without ctenidium at hypandrium (Suppl. material |
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– | Male genitalia always with ctenidium at hypandrium (as in Suppl. material |
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5 | Large species (15–20 mm) with long body pilosity and broad metafemur covered with long pile (Suppl. material |
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– | Species with shorter pilosity and different combinations of characters |
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6 | Sternum 4 medially clearly divided with membranous structure and with posterolateral tubercles or laminate extensions (Suppl. material |
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– | Sternum 4 and basotarsomere of metatarsus without such modifications |
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7 | Male genitalia: posterior surstyle lobe divided into two branches (Suppl. material |
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– | Male genitalia: posterior surstyle lobe not divided into branches; eyes holoptic |
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8 | Abdomen elongated and narrow; terga black; terga 2–4 with a pair of pollinose fasciate maculae (Suppl. material |
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– | Species with different combinations of characters |
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9 | Basoflagellomere elongated, at least three times as long as wide (Suppl. material |
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– | Basoflagellomere less elongated; posterior surstyle lobe different |
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10 | Posterior surstyle lobe with basolateral protrusion (lateral hump) (Suppl. material |
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– | Posterior surstyle lobe of male genitalia without basolateral protrusion (lateral hump) |
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11 | Basoflagellomere reddish-yellow (Suppl. material |
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– | Basoflagellomere brown to black; tarsi of metaleg dark; metatarsus shorter, two times longer than wide (Suppl. material |
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12 | Basoflagellomere with convex dorsal margin (Suppl. material |
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– | Basoflagellomere with concave dorsal margin (Suppl. material |
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13 | Metafemur more incrassate, ca. three times longer than wide, covered with dense pilosity (Suppl. material |
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– | Metafemur less incrassate, at least three times longer than wide (as in Suppl. material |
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14 | Basoflagellomere shorter, 1.3 times as long as wide, with convex dorsal margin (Suppl. material |
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– | Basoflagellomere elongated, two times as long as wide (Suppl. material |
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15 | Small sized species (8-11 mm) with metallic shiny body; scutum and terga strongly punctate, without or with very weak pollinose areas (Suppl. material |
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– | Species with different combinations of characters |
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16 | Metafemur with short pilosity (Suppl. material |
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– | Metafemur with longer pile (as in Suppl. material |
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17 | Abdomen broad, oval (Suppl. material |
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– | Species with different combinations of characters |
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18 | Basotarsomere of metatarsus elongated, four times as long as wide (Suppl. material |
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– | Species with different combinations of characters |
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19 | Abdomen narrow, elongated (Suppl. material |
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– | Species with broader abdomen and metaleg |
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20 | Tergum 2 without or with indistinct narrow pollinose fasciate maculae (Suppl. material |
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– | Tergum 2 with broad pollinose fasciate maculae (Suppl. material |
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21 | Terga 2–4 strongly punctate; second and third tarsomeres similar in size (Suppl. material |
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– | Terga 2–4 finely punctate; second tarsomere longer than third (Suppl. material |
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22 | Females (genitalia not visible) |
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– | Males (genitalia visible externally) |
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23 | Metatibia swollen in apical half (Suppl. material |
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– | Metaleg without such modifications |
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24 | Posterior surstyle lobe with basolateral protrusion (lateral hump) (Suppl. material |
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– | Posterior surstyle lobe without basolateral protrusion |
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25 | Face with a bulge below antennae (Suppl. material |
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– | Face without a bulge below antennae |
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26 | Metatrochanter without calcar |
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– | Metatrochanter with distinct calcar (Suppl. material |
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27 | Basoflagellomere 1.2 times as long as wide (Suppl. material |
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– | Basoflagellomere short, as long as wide (Suppl. material |
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28 | Basoflagellomere elongated, at least three times as long as wide (Suppl. material |
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– | Basoflagellomere shorter, less than three times as long as wide (as in Suppl. material |
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29 | Eye contiguity very short, approximately four to five facets long (Suppl. material |
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– | Eye contiguity more than 10 facets long (as in Suppl. material |
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30 | Tarsi yellow dorsally and ventrally (Suppl. material |
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– | Tarsi dark brown/black dorsally and orange/brown ventrally (Suppl. material |
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31 | At least terga 2 and 3 with brown, reddish or yellow markings |
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– | Only tergum 2 with brown, reddish or yellow maculae, other terga dark |
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32 | Metatibia swollen in apical half (Suppl. material |
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– | Metatibia of normal shape (as in Suppl. material |
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33 | Pile on ventral margin of metafemur dense and long, the longest as long as width of metafemur (Suppl. material |
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– | Pile on ventral margin of metafemur shorter, maximum as long as half of width of metafemur (as in Suppl. material |
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34 | Basoflagellomere shorter, 1.3 times as long as wide, with convex dorsal margin (Suppl. material |
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– | Basoflagellomere longer, with straight or concave dorsal margin (Suppl. material |
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35 | Tarsi yellow, tibiae mostly yellowish, only medially brown; frons and vertex usually partly reddish to yellow (Suppl. material |
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– | Legs mostly black, at least tarsi dark; frons black |
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36 | Basoflagellomere elongated, more than 1.5 times as long as wide (Suppl. material |
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– | Basoflagellomere shorter, less than 1.3 times as long as wide (Suppl. material |
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37 | Basoflagellomere very short, as long as wide (Suppl. material |
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– | Basoflagellomere longer, 1.2 times as long as wide (Suppl. material |
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38 | Face with a bulge below antennae (Suppl. material |
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– | Face without a bulge below antennae |
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39 | Basoflagellomere elongated, at least 2.7 times as long as wide (Suppl. material |
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– | Basoflagellomere shorter, less than 2.5 times as long as wide (as in Suppl. material |
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40 | Tarsi yellow dorsally and ventrally (as in Suppl. material |
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– | Tarsi dark brown/black dorsally and orange/brown ventrally (as in Suppl. material |
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1 | Oral margin reduced, covered by microtrichia (Suppl. material |
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– | Oral margin notched, slightly produced forward (as in Suppl. material |
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2 | Metatrochanter with sparse pale pile (Suppl. material |
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– | Metatrochanter with dense and strong yellow to red brush of pile (Suppl. material |
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3 | Apical fourth of tibiae and all tarsi bright yellow; Afrotropical species (Zimbabwe) |
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– | Tarsi partly brown or black; Palaearctic species |
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1 | Scutum with distinct pollinose ornamentation, vittae and fasciae (Suppl. material |
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– | Scutum with indistinct pollinose vittae (Suppl. material |
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The
The
The
The
The
Out of 194 described species (234 in total including undescribed taxa), 180 (209) species are distributed in the Palaearctic Region and 14 (27) are known from the Afrotropical Region. Three lineages (
The
The
The
The
The
At present and based on our results, the regions with the highest species richness are the Mediterranean Peninsulas: Iberian, Balkan and especially Anatolian. Certain areas in the Palaearctic (regions of Pakistan, Central Asia and eastern part of the Middle East) and Afrotropical Regions (Central and Eastern Africa) have been under-sampled and they need additional collecting efforts. Central Asia and Pakistan are characterised by numerous endemics with potential significance to understand the evolutionary scenario of the genus
We thank Branka Božičić Lothrop for valuable comments and editing of the manuscript, and Hugh Lothrop, native English speaker, for linguistic revision. The study was supported by the Ministry of Education, Science and Technological Development, Republic of Serbia, Grant No. 173002, III43002, 451–03–68/2020–14/200358 and 451–03– 68/2020–14/200125, the Provincial Secretariat for Science and Technological Development of the Republic of Serbia (Grant No. 114–451–1125/2014–03 and 114–451–1702/2014–03).
Figures S1–S37: Figures of morphological characters
multimedia
Figures S1–S13: Figures of male genitalia
multimedia
Figures S1–S15: Figures of adults
Adobe PDF file
List of
species data
Table S1. Classification of the genus
species data