Corresponding author: Christian Schmid-Egger (
Academic editor: A. Köhler
The genus
Schmid-Egger C, van Achterberg K, Neumeyer R, Morinière J, Schmidt S (2017) Revision of the West Palaearctic
The paper wasp genus
The taxonomy of the
The present study employs the concept of integrative taxonomy (
Specimens for DNA barcoding are primarily deposited in the collections of the Zoologische Staatssammlung München and the private collection of
For the present study, a large number of specimens from several collections were morphologically examined. The taxonomic treatment of species is primarily based on the combined analysis of morphological and molecular data, and only those specimens that were analysed both, morphologically and genetically, are listed in the Suppl. material
DNA extraction, PCR amplification, and sequencing were conducted at the Canadian Centre for DNA Barcoding (
Sequence divergence statistics were calculated using the Kimura two-parameter model of sequence evolution (
Barcoding statistics of
BIN | n | Country | Mean intraspecific distance | Maximum intraspecific distance | Nearest |
Distance to NN | |
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12 | CH, DE, KZ | 0.03 | 0.16 |
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2.31 |
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7 | HR, IT | 0 | 0 |
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4.82 |
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2 | IT | 0 | 0 |
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1.7 |
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5 | CH, FR, MA | 0.58 | 0.95 |
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2.18 |
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21 | DE, IT | 0.57 | 1.17 |
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2.48 |
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11 | CH, FR, HR, SP | 0.14 | 0.32 |
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2.31 |
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9 | AZ, CY | 1.03 | 2.03 |
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3.29 |
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1 | AZ | |||||
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3 | GR | |||||
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14 | DE, FR | 2.45 | 5.42 |
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3.29 |
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37 | AZ, CH, DE, GR, HR, IT | |||||
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10 | MA | |||||
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21 | AZ, GR, HR, IT, TR | 0.49 | 1.29 |
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2.88 |
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20 | HR, IT, PT, SP | 0.28 | 1.09 |
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2.49 |
NONE | 2 | AZ, DE | |||||
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1 | MA | N/A | 0 |
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3.78 |
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51 | AZ, CY, GR, HR, IT, TR | 0.88 | 2.04 |
|
2.99 |
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18 | DE, GR, IT | 2.2 | 6.28 |
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3.56 |
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4 | DE | |||||
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6 | CH, IT | 0.28 | 0.65 |
|
1.7 |
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4 | UAE | 0 | 0 |
|
12.84 |
The
Regional terms of geographic distribution are abbreviated as Central (C), Southern (S), Eastern (E), Northern (N) and West (W). Southwest Asia includes Turkey, Iran, Israel and adjacent countries. Temperate Asia refers to Asia north of 37° northern latitude (approximative only, species may occur south of 37° north, e.g. in mountain regions).
1 | Parastigma elongate and long compared to length of vein 1-SR of fore wing (a); second and third metasomal tergites with brown or blackish curved lines (b); [mesopleuron only sparsely punctate medially]; |
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– | Parastigma short, hardly or not longer than wide and short compared to length of vein 1-SR of fore wing (aa); second and third metasomal tergites without curved lines, often with a black pattern (bb) |
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2 | Female: Dorsal part of epistomal (frontoclypeal) suture blackish or dark brown (a); length of fore wing 15–28 mm. Male: Clypeus evenly convex (b); lateral tubercles on each side of apex of last sternite subtriangular and wider basally (c), its terminal apophyses long and spatulate apically (d); Palaearctic distribution: Egypt, Oman, Iran, Afghanistan |
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– | Female: Dorsal part of epistomal suture yellowish (aa); length of fore wing 11–17 mm. Male: Clypeus with impression (bb); lateral tubercles of last sternite more cylindrical and narrower basally (cc), its terminal apophyses long and pointed apically (dd); Palaearctic distribution: Afghanistan, Iran, Iraq, Saudi Arabia, United Arab Emirates, Oman, China |
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3 | Body brown with yellow pattern (a); mesopleuron indistinctly sculptured (b, c); recently introduced to northern Spain, native to South and Central America, Caribbean islands and southern U.S.A. ( |
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– | Body black with yellow pattern (aa); mesopleuron with dense irregular microstriae (bb, cc) ( |
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4 | Antenna with 12 segments (a); metasoma with 6 visible tergites and sternites (b); face and/or clypeus with yellow and black pattern (c); females |
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– | Antenna with 13 segments (aa); metasoma with 7 visible tergites and sternites (bb); face and/or clypeus yellow (cc; but more or less blackish pattern present in |
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5 | Mandible very stout and its outer face more or less depressed (a); ventral border of clypeus strongly convex (b) and medially acute (c); socio-parasitic species |
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– | Mandible comparatively slender and its outer face weakly convex or flat (aa); ventral border of clypeus flat (bb) and medially truncate or slightly concave (cc); social species |
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6 | Basal half of mandible distinctly angulate (a) and flattened (b); clypeus gradually depressed ventrally (c); [yellow area along inner eye margin connected to yellow bar above antennal sockets; clypeus largely punctate ventrally and with fine pubescence as in medial area (c)]; SE and southern C Europe, C Asia [= |
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– | Basal half of mandible gradually curved (aa) and convex (bb); clypeus abruptly depressed ventrally (cc) |
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7 | Dorsal ridge of mandible wide, convex and distinctly elevated above middle of mandible (a, b); fine pubescence of clypeus conspicuous and comparatively long (b, c); yellow area along inner eye margin usually connected with yellow bar above antennal sockets (d); SW and southern C Europe, N Africa; [= |
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– | Dorsal ridge of mandible narrow, slightly convex and hardly elevated above middle of mandible (aa); fine pubescence of clypeus inconspicuous and short (bb); yellow area along inner eye margin separated from yellow bar above antennal sockets (cc) |
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8 | Clypeus 0.7 times as long as wide (a); lower ridge of mandible 0.4 times as wide as mandible (b); depression of mandible with rather dull surface (c); area above lower edge of clypeus (below dark spot) dull and straight (d); S and southern C Europe to W Asia |
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– | Clypeus 0.8–0.9 times as long as wide (aa); lower ridge of mandible 0.2 times as wide as mandible (bb); depression of mandible with shiny surface (cc); area above lower edge of clypeus (below dark spot) shiny and distinctly convex (dd); Morocco |
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9 | Malar space mainly yellow and connected to yellow ventral part of temple (a); mandible usually black (b), |
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– | Malar space black as ventral part of temple (aa); |
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10 | Specimens from Europe (excluding Crete) and NW Africa |
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– | Specimens from Asia (including Crete and Cyprus) |
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11 | Transverse yellow band of pronotum narrower medio-laterally than dorso-laterally near junction with oblique yellow stripes of pronotum (a), rarely narrowed dorso-laterally and slightly widened below it |
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– | Transverse yellow band of pronotum as wide medio-laterally as dorsolaterally or wider (aa; rarely narrowed medio-laterally and slightly widened below it) |
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12 | Scapus slightly widened apically in dorsal view (a), if scapus intermediate then ocellar triangle acute anteriorly (b); change in sculpture between mesepisternum and epicnemium frequently gradual (c; = epicnemial ridge indistinct); apical half of antenna more or less brownish dorsally (d); apical half of hypopygium (= sternite VI) often largely black and brown or largely brown, slightly darker than apex of last tergite (e); border of black and yellow on outer side of middle and hind femora often sharp and often without orange intermediate area in Balkan populations (f); often smaller species; S Europe, W Asia |
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– | Scapus distinctly widened apically in dorsal view (aa); ocellar triangle transverse (bb); change in sculpture between mesepisternum and epicnemium abrupt (cc; = epicnemial ridge distinct); apical half of antenna orange or yellow (dd); apical half of hypopygium yellow or largely so and as pale as apex of apical tergite (ee), rarely darker; border of black and yellow on outer side of middle and hind femora washed out, partly because of orange intermediate area (ff); somewhat larger species); Europe except N, NW Africa, W and C Turkey, Azerbaijan, probably also farther east in temperate Asia. Not in Crete, see |
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13 | Apical half of sternite VI yellow or largely so, and as yellow as apex of last tergite (a); ventral part of mesopleuron in general more coarsely sculptured than remaining parts (b); [mesoscutum usually with two yellow spots; apical half of antenna orange or yellow reddish] |
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– | Apical half of sternite VI largely black, |
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14 | Temple (gena) with wide yellow band, medially wider than half width of temple (a); clypeus yellow medially (b); propodeum and second tergite largely yellow (c); incision of eye mainly yellow (d); [frons slightly convex (about similar to area between antennal sockets and distinctly less so in |
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– | Temple with separated yellow spots (aa), |
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15 | Posterolateral yellow bands of pronotum usually not connected to anterior transverse band (a); anterior transverse yellow band of pronotum medio-laterally as wide as dorso-laterally or wider (b); mesoscutum with paired yellow spots (c); [clypeus with transverse black band; hypopygium all black, with apical yellow or reddish spot or entirely reddish; distinction from |
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– | Posterolateral yellow bands of pronotum usually connected to anterior transverse band (aa); anterior transverse yellow band of pronotum medio-laterally narrower than dorso-laterally (bb); mesoscutum sometimes lacking yellow spots (cc); [colour pattern is highly variable in this species, more so than in Europe]; Europe and Palaearctic Asia |
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16 | Transverse yellow band of pronotum narrow (a); antenna largely black or blackish dorsally (b), similar to colour of frons; basal third of scapus more or less black or blackish ventrally (c); pubescence of pronotum and mesoscutum 0.7–0.9 times as long as diameter of anterior ocellus (d); yellow spots of propodeum narrow or absent, |
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- | Transverse yellow band of pronotum wider (aa); antenna orange brown or somewhat darkened dorsally (bb) and distinctly paler than frons; basal third of scapus yellow ventrally (cc), but darkened in most |
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17 | Malar space 1.22–1.76 times as long as |
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– | Malar space 0.87–1.19 times as long as |
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18 | Malar space 0.85–1.0 times as long as |
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– | Malar space at most 0.75 times as long as |
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19 | Apical yellow band of sternite IV medially interrupted (a) or narrow; hind coxa black dorsally (b); clypeus often with a wide black transverse band or with more or less transverse trapezoid black patch, often close to or connected with lateral margins of clypeus and situated nearly halfway clypeus (c); change in sculpture between mesepisternum and epicnemium frequently gradual (d; = epicnemial ridge indistinct); southern C and S Europe |
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– | Apical yellow band of sternite IV usually complete and wide (aa); hind coxa with yellow patch dorsally (bb); central dark patch on clypeus usually less developed and situated below middle of clypeus (ccc) or absent (cc); usually with an abrupt change in sculpture between mesepisternum and epicnemium (dd; = epicnemial ridge distinct), but sometimes rather gradual |
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20 | Transverse yellow band of pronotum medio-laterally wider than dorso-laterally (a); central dark patch on clypeus usually developed, but small, rounded or forming a transverse band (b); mesoscutum usually with pair of medium-sized to large yellow spots (c); [some specimens have an all yellow clypeal disk and all black mesoscutum; specimens from NW Africa often have sternite IV apically yellow, which is more or less darkened in northern specimens]; NW Africa, SW Europe, N Italy, southern Switzerland, Croatia, Corfu |
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– | Transverse yellow band of pronotum dorso-laterally wider than medio-laterally (aa); clypeus entirely yellow (bb) or with minute black spot, but spot sometimes medium-sized (bbb); mesoscutum usually black (cc) or with pair of small yellow spots (but sometimes large in Asian specimens; ccc); Croatia and SE Europe, Turkey, Cyprus to Central Asia and Egypt |
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21 | Mandible very stout and with a distinct depression on its outer face (a); clypeus depressed medio-apically (b), its margin tapered to a small point (c); malar space comparatively long and wide (d); socio-parasitic species |
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– | Mandible comparatively slender and its outer face flat or slightly convex (aa); clypeus flat medio-ventrally (bb), its margin evenly convex or broadly triangular (cc); malar space medium-sized (dd) or narrow; social species |
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22 | Basal half of mandible sinuate in dorsal view (a); mandibular depression strongly concave (b); [mandible mainly yellow or brown (d); fore and middle coxae and mesopleuron ventrally usually with yellow pattern (c); dorsal ridge of mandible at most 0.25 times mandibular width and prominent]; SE and southern C Europe to Central Asia |
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– | Basal half of mandible straight in dorsal view or nearly so (aa); mandibular depression shallowly concave (bb); [fore and middle coxae and mesopleuron ventrally usually black (cc), colour of mandible variable (d, dd)] |
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23 | Mandible mainly yellow, except for its more or less darkened margins (a); mandibular depression shorter and occupying less than half of outer face of mandible (b), dorsal ridge at least 0.33 times as wide as mandible (c); clypeus entirely yellow medially (d); SW Europe, southern C Europe, NW Africa |
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– | Mandible black (aa), rarely with small yellow spot; mandibular depression wide and occupying most of outer face of mandible (bb), dorsal ridge 0.25–0.30 times mandibular width (cc); clypeus medially with dark brown pattern (dd), but sometimes largely reduced or absent; S Europe and southern C Europe to W Asia. [If from Morocco consider |
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24 | Temple bulged behind eye in dorsal view, slightly convex (a); head trapezoid in anterior view (b); apical margin of clypeus triangular (c); width of clypeus 1.0–1.1 times its length (d, ddd); latero-ventral margin of clypeus narrowly black or dark brown and convex in lateral view (e), but flattened in |
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– | Temple narrowed behind eye in dorsal view, more or less straight (aa); head nearly triangular in anterior view (bb), but less so in |
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25 | Dorsal length of apical antennal segment 2.2–2.9 times as long as its maximal width (a), 1.3–1.5 times as long as fifth antennal segment (b); clypeus distinctly depressed medially (c) and with distinct lateral ridges (d); medio-longitudinal depression of face more or less impressed and U-shaped (e); antenna coloured according to fig. f; short carina between antennal sockets sharp dorsally and usually pale yellow (g); clypeus with short bristles and medio-ventrally flat; width of clypeus 1.0–1.1 times its length; [head sometimes distinctly narrowed ventrally]; Europe and Palaearctic Asia |
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– | Dorsal length of apical antennal segment 1.5–2.1 times as long as its maximal width (aa), 1.0–1.2 times as long as length of fifth antennal segment (bb); clypeus not or slightly depressed medially (cc) and lateral ridges absent dorsally or slightly developed (dd); face flat medially, without longitudinal depression (ee); antenna coloured as in ff or fff; short carina between antennal sockets obtuse dorsally and more or less infuscate (gg); width of clypeus 1.1–1.2 times its length |
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26 | Mesosternum entirely black (a); black baso-dorsal stripe on third antennal segment 0.25 times length of segment (b); lateral ridges of clypeus and ridge between antennal socket and clypeus more distinct (c); [largely yellow coloured species, specimens from eastern Turkey however may have reduced whitish yellow markings; W and C Asia, Cyprus, Egypt, Crete. |
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– | Mesosternum at least with some yellow spots posteriorly (aa), and often entirely yellow, but in some males of |
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27 | Most of clypeus with long bristles and distinct punctures (a); apical half of antenna uniformly orange-yellow dorsally and ventrally (b); apical antennal segment comparatively stout and dorsally about 1.5 times as long as wide basally (c); setae of pronotum medio-dorsally and of mesoscutum about half as long as width of posterior ocellus (d); width of clypeus 1.1 times its length (e) and medio-ventrally more flattened (f); [fourth and fifth antennal segments strongly oblique in lateral view]; Europe (for specimens from Crete see also |
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– | Dorsal 0.7 of clypeus without bristles and punctures (aa); apical half of antenna usually darker dorsally than ventrally (bb); apical antennal segment more slender, dorsally about twice as long as wide basally (cc); setae of pronotum medio-dorsally and of mesoscutum at least 0.8 times as long as width of posterior ocellus (dd) or longer; width of clypeus about equal to its length (ee) and medio-ventrally less flattened (ff); [fourth and fifth antennal segments moderately oblique in lateral view]; Europe, temperate Asia |
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28 | Ventral half of clypeus distinctly depressed (a) and with distinct lateral ridges (b); dorsal length of apical antennal segment about 3.0 times its width (c); frons with a distinct longitudinal depression medially (d); sternites III-VII usually with transverse yellow band basally (e; visible with sternites sufficiently extruded) and last sternite often partly yellow (f); apical half of antenna usually distinctly darkened dorsally (g); clypeus more or less truncate medio-ventrally (h); S Europe, W Asia |
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– | Ventral half of clypeus at most slightly depressed (aa) but without lateral ridges (bb); dorsal length of apical antennal segment about twice its width (cc), but 2.2–2.7 times in |
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29 | Malar space long, 0.8–1.0 times as long as width of third antennal segment measured basally (a, b); malar space often entirely yellow (c) or narrowly black posteriorly; [basal half of second sternite with yellow pattern; mesopleuron obliquely rugulose (d)] |
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– | Malar space short, 0.4–0.6 times basal width of third antennal segment (aa, bb); malar space black posteriorly with black area often widened (cc) |
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30 | Antenna dorsally orange-yellow (a); pronotal transverse yellow band more or less widened (b); epicnemial ridge delicate but well-defined (c), sometimes reduced in Peloponnesian populations; mesoscutum almost always with pair of medium-sized to large yellow spots (d), sometimes minute or absent; scutellar spots large (e); inner side of hind tibia often partly darkened (f); medial area of face narrower; N Italy to SE Europe, Turkey and Central Asia |
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– | Antenna dorsally dark brown (aa); pronotal transverse yellow band narrow (bb); epicnemial ridge indistinct (cc); mesoscutum with yellow spots minute or absent (dd); scutellar spots small (ee); inner side of hind tibia rather pale (ff); medial area of face wider; C Europe, temperate Asia |
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31 | Specimens from W to C Asia, Cyprus, Egypt |
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– | Specimens from Europe and NW Africa |
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32 | Middle and hind coxae largely black dorsally (a); apical yellow band of sternite VI medially interrupted (b); Turkey |
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– | Middle and hind coxae largely yellow dorsally (aa); apical yellow band of sternite VI medially continuous (bb); Cyprus, Turkey to Central Asia and Egypt |
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33 | Mesosternum entirely black or with a pair of elongate yellow spots (a); basal half of second metasomal sternite entirely black (b); oblique yellow stripes of pronotum often long or medium-sized (c); frontal ridge narrower (d); apical antennal segment slightly slenderer and usually parallel-sided basally (e); SE Europe |
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– | Mesosternum predominantly or entirely yellow (aa); basal half of second sternite with yellow pattern, varying from a pair of very small sublateral yellow spots to mainly yellow (bb), |
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34 | Yellow transverse pronotal band narrow laterally (a), |
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– | Yellow transverse pronotal band moderately to strongly widened laterally, of even width medio-laterally (aa); apical half of antenna uniformly orange to yellow dorsally and ventrally (bb), mesoscutum nearly always with minute to large paired yellow spots (cc); scutellum with large yellow spots (dd); setae of pronotum and mesoscutum usually shorter, about 0.5 times diameter of posterior ocellus and mainly straight (ee); NW Africa, SW Europe, N Italy, S Switzerland, Croatia, Corfu |
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The species typically occurs on humid meadows or along lake shores or in fens with a large reed zone, but it also colonizes dry habitats. In contrast to
The female is characterized by reduced yellow markings and dorsally dark antennal segments. It can be confused mainly with
The male is unique within the
Europe: France (eastern part), Belgium, Switzerland, Germany, Austria, Czech Republic, Russia (Orenburg Oblast). Asia: Kazakhstan, Mongolia, Russia (Primorsky Krai), China.
Specimens from Switzerland, Germany, and regions as far apart as Kazakhstan exhibited little intraspecific variation (0.16%, Table
The recognition of
The male is unique by the combination of narrow temples (genae) in dorsal view and a markedly depressed clypeus with distinct lateral ridges. The dorsal length of the apical antennal segment is about 3.0 times its maximum width, and longer than in similar species.
Southern Europe and Turkey, northwards to Switzerland, southwards to Israel, eastwards to Azerbaijan.
Europe: Spain, France, Italy, Switzerland, Croatia, Bulgaria, Macedonia, Montenegro, Greece. Asia: Israel.
We regard
The social parasitic species can be recognized by the shape of the mandibular impression and by the colour pattern of the clypeus. Females of the species group show differences in the depth of the medial impression of the mandible and the size and shape of the upper ridge. The weakest medial impression occurs in
The shape of the mandibular impression in males generally follows that of females, but is in general less developed. In addition, the male of
Southern C and S Europe, northwards to S Germany, Turkey, Iran, Armenia (Guiglia, 1972). Records from NW Africa probably refer to
Europe: Bulgaria (Rhodope Mts), France, Greece, Italy (Alps, Abruzzi), Spain, Switzerland.
Not enough specimens were sequenced to detect any genetic variation.
Holotype, ♀ (
Konstantin Samartsev (
Large and relatively bright species with robust mandible and wide, yellow malar space (Figs
FEMALE. Holotype, body length 15.8 mm; fore wing length 11.6 mm.
For colour pattern, see figures.
MALE. Body length 11.6–17.1 mm; fore wing length 9.6–11.7 mm. For colour pattern, see figures. Similar to female, differs as follows:
Specimens from Algeria, Andorra, France, Italy, Morocco, Portugal, Spain, and Switzerland have been examined, indicating that the species is confined to NW Africa and SW Europe with an extension to Central Europe, and is replaced by
According to
Specimens from south-central Europe and Morocco only showed a small genetic distance.
The name is a combination of the Latin adjectives “australis” (southern) and “occidentalis” (western), because of its southwestern distribution in Europe.
See
The female shares a dark brown to black upper side of antennal segments with
Europe including Norway and Sweden south of 65° N to Turkey, Central Asia.
Europe: Austria, Belgium, France, Germany, Greece, Italy, Netherlands, Sweden, Switzerland. Asia: Tajikistan, Turkey, Uzbekistan.
For a detailed discussion about the mixed type series of the species see
This species belongs to the group of species with a short female malar space within the
S Europe and Turkey, northwards to Austria (Neusiedl am See) and Switzerland (northwards to Zürich). See
Europe: Spain, Gibraltar, Switzerland, southern France and Corsica, Austria, Greece, Italy, Croatia. Asia: Eastern Turkey (Hakkari region).
Specimens from Spain, S France and Corsica, Switzerland, and Croatia were barcoded. The species forms a separate cluster, with a small gap between SW (France, Spain) and southern central European populations.
Specimens from Mongolia and China (=
The species was first described from Uzbekistan and later as
Another problematic taxon in this group is
According to the description by
Based on the material that was available for us it appears that two taxa of this lineage occur in eastern Turkey and the Caucasus:
The
Three barcoded specimens from Crete were assigned a separate BIN (Tab.
The specimens barcoded fall into four clusters that were assigned three different BINs (Suppl. material
A single specimen from Azerbaijan has been assigned a different BIN, whereas another specimen from Azerbaijan that is morphologically similar to the remaining females of
Females of
Sternite II is predominantly black with narrow apical yellow band in
Males of
Females from Crete (n = 6) differ from typical
The single male that was examined genetically has the mesosternum all black and the yellow band of the temple is medially interrupted. In specimens from Iran the yellow body colour of the only examined male from Arak is replaced by an extreme whitish yellow. For recognition of specimens from Caucasus and E Turkey see discussion on
From Central Turkey to Central Asia, Israel, Iran and Egypt. In Europe only known from Crete and Cyprus. Specimens described from China and Mongolia (not examined) may also belong to
Europe: Greece (Crete), Cyprus. Asia: Turkey (Diyarbakir, Hakkari, Van, Esendere, Mersin, Göreme, Kars/Kagisman), Iran (Arak/Besril), Azerbaijan, Uzbekistan (Bukhara), Israel (Arava Valley). Africa: Egypt (female, Oasis Dakhla, 2.ix.1992,
Phylogenetic tree resulting from Bayesian analysis of
Two of these clusters (2 and 3) occur in Europe. The current data indicate for cluster 2 a south-western European distribution although we could only examine one specimen from France and records from Spain still are missing. The cluster occurs over whole Germany and it is close to the cluster of Morocco (1). The other European cluster (3) includes specimens from Greece and Azerbaijan and seems to have a more south-eastern distribution. It covers whole Germany with the westernmost records from the Aosta valley in the Italian Alps. This cluster is genetically closer to
The high intraspecific variation (Table
The population of Crete seems to be isolated from the remaining populations for a long period, and the common ancestor probably came from the
Because of this result, we follow the concept of morphospecies here and treat
The female of
The male of
Some specimens of
NW Africa, C and S Europe as far north as Latvia, but missing in Great Britain, Scandinavia, Crete and Cyprus. Introduced to Australia, North America (including Canada,
Europe: Examined from most countries in Central and S Europe. Asia: Turkey (Termessos/Antalya), Azerbaijan. Africa: Morocco, Tunisia.
The species is widespread in N Italy, the Balkans, and western Asia (Turkey to Caucasus area). The identity of
From NE Italy to Greece and Azerbaijan. Widespread and common in mainland Greece (
Europe: Italy (Trentino/Rovereto, Lombardia/Valtellina/Grossio, Veneto/N of Verona), Croatia (Krk), Greece (Crete/Matala, Peloponnese), Cyprus (Akrotiri). Asia: Turkey (Antalya), Azerbaijan.
Specimens of
The name
Western and central Mediterranean area, eastwards to the Greek island of Corfu. In NW Africa from Tunisia to Morocco. Northwards to Italian Alps and S Switzerland. Specimens from Egypt and Israel formerly assigned to
Europe: Croatia (Istria), Greece (Corfu), Italy (Lombardia/Brescia, Veneto/N. of Verona, Pavone, Dro, Sardinia), Spain (Mallorca/Alcúdia, Andalucía/various locations), France (Bouches du Rhône/Alpilles, Pont du Diable, La Rouquette), Portugal (Algarve/Carrapeteira), Malta (Ghajn Tuffieha, female, photo from Kristofer Mogyorossy), Switzerland. Africa: Algeria (Alger), Morocco (Sefrou, Ht Atlas), Tunisia (Le Kef).
Specimens from
Holotype♀, Morocco, Haut Atlas, 2 km N Tizi n’Tichka,
FEMALE. Holotype, body length 14 mm; fore wing length 11.5 mm.
For colour pattern see figures.
Characters for separating
Lower ridge of mandible narrow, in HT= 0.22× (in PT= 0.24×) as large as mandible (measured medially) | Lower ridge of mandible wider, 0.37× as wide as mandible |
Both ridges of clypeus smaller, upper ridge in PT nearly flat and barely expressed | Both ridges of clypeus wider |
Medial impression of mandible shiny, with very fine microsculpture | Medial impression of mandible dull, with coarser microsculpture |
Lower margin of clypeus (= near margin of yellow colour) ridged, shiny | Lower margin of clypeus straight, with microsculpture |
Mesoscutum in HT medially with two minute yellow spots, black in PT | Mesoscutum all black |
Black clypeal spot band-shaped, small in holotype, missing in paratypes | Black clypeal spot larger |
Only known from the High and Middle Atlas Mountains in Morocco. Previous to this study,
The species is most probably a social parasite. At the type locality, it was collected together with
DNA barcoding of a specimen from Morocco, formerly identified as
The species is widespread in SE Europe to C Asia and China. Apart from the original description it was later described as
Within the
The recognition of
In N Africa
Balkans from Croatia to Greece, east to Central Asia, Mongolia, and China, south to Israel and Egypt.
Europe: Croatia, Serbia, Macedonia, Greece, Cyprus. Asia: Turkey (Antalya, Hakkari), Israel (Jordan Valley), Azerbaijan, China. Africa: Egypt (Kairo; Al Fajum).
Specimens from Croatia, Greece, Turkey, Cyprus, and Azerbaijan were analysed.
Differences in the ocellar angle (more obtuse in
The most important diagnostic character of
The latter character is used here for recognition of
The separation of
Europe, north to S Finland, Palaearctic Asia east to Mongolia, China, and Russian Far East.
Europe: Germany, Italy (Alps), Bulgaria, Greece, France, Croatia, Portugal, Switzerland. Asia: Iraq, Turkey (Kars/Kagisman, Hakkari region, Denizli/Pamukkale, Antakya, Alanya, Marmaris, Diyarbakir).
Only specimens from Central Europe were examined genetically, except for one specimen from Greece. The species shows significant intraspecific genetic variation. It is possible that the examination of Asian species will yield unexpected results.
The species was formerly treated as
The species can only be recognized based on the shape of the mandible, clypeus, and the colour of male fore and mid coxae. The upper ridge of the mandible is markedly modified in the female and forms a triangle in dorsal view (weaker and more rounded in male). The recognition of males is more difficult because the upper ridge is sometimes only weakly curved and resembles that of
S and C Europe, north to Germany (Hesse, one record from 1908;
Europe: France, Croatia, Italy (Alps, Abruzzi, Calabria, Sicily), Greece, Macedonia, Montenegro, Serbia, Switzerland. Asia: Azerbaijan, Turkey (Uludağ, Bursa, Van), Syria, Egypt, Turkmenistan.
Barcoded specimens from south-central Europe showed little genetic variation.
The following species are only discussed briefly, because they were either introduced only recently or they occur near the southern border of the study area. For further information on the nomenclature of these species see
The nominate subspecies of
The species is recorded from E Africa, S Asia and Australia (
The species is recorded from SW and S Asia (Arabian Peninsula, Iran, India, and S China) according to
The phylogenetic analysis based on Bayesian inference resulted in a split of the included
These results concur with an earlier analysis of a mitochondrial gene fragment (16S ribosomal RNA) by
The most important change affected the two species
Species and species groups of
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Likewise,
Despite the comparatively limited informative value of a single mitochondrial gene region like the 658bp
We are greatly indebted to Khalid A. Aliyew (National Academy of Sciences of Azerbaijan, Baku) for loaning us very important material (
List of voucher specimens
Microsoft Excel Worksheet (.xlsx)
specimens date.
NJ tree
Adobe Acrobat Document (.pdf)
phylogenic data.