Corresponding author: Cornelis van Achterberg (
Academic editor: Andreas Köhler
The European species of the potter wasp genus
van Achterberg C, Smit JT, Ljubomirov T (2023) Review of the European
The potter wasp genus
The setosity of body parts is used extensively in existing literature. However, this is a variable and, therefore, rather problematic character, augmented by wear of the setae in aged specimens and concealed setae in wet and/or dirty specimens. In one species (
Females build nests consisting of one or several jug like mud cells (Fig.
Identified material was used from the following collections:
Showing dark (
DNA extraction was conducted on single legs, using the NucleoMag 96 Tissue kit by Macherey-Nagel on a Thermo Scientific KingFisher Flex magnetic bead extraction robot, with a final elution volume of 150 µl. The standard COI barcoding fragment (
Sampled specimens from
Taxon | ID number | BOLD accession number | Country |
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Netherlands | |
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Netherlands | ||
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Netherlands | |
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Netherlands | |
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Netherlands | |
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Netherlands | |
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Netherlands | |
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Netherlands | |
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Bulgaria | |
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Bulgaria | |
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Bulgaria | |
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Bulgaria | |
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Bulgaria | |
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Bulgaria | |
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Bulgaria | |
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Netherlands | |
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France | |
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Greece | |
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Bulgaria | |
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Bulgaria | |
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Greece | |
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Finland |
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Finland |
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Netherlands |
In the Neighbour-Joining tree (using COI sequences) newly barcoded specimens from Bulgaria, France, Greece, and Netherlands are combined with unpublished older sequences from Finland and Netherlands (Fig.
First of all, it is clear that
Neighbour-Joining tree for barcoded European
The position of the
N.B. Species can only be reliably identified by a combination of characters. Setosity may be worn off in aged specimens or hardly visible in dirty specimens; therefore, aged or dirty specimens are easily misidentified. In addition, there is a considerable intraspecific variation.
1 | Females: antenna without terminal hook (a); clypeus often partly or entirely black (b); inner tooth of hind tarsal claw widened and apically more or less truncate (c), but intermediate in |
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– | Males: antenna with terminal hook (aa); clypeus nearly always entirely yellow (bb); inner tooth of hind tarsal claw comparatively slender and apically acute (cc) |
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2 | Setae of occiput short to medium-sized (a); second and third antennal segments reddish or brownish ventrally (b), rarely entirely black in SW Europe; [pronotum often broadly yellow laterally] |
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– | Setae of occiput long (aa; but sometimes worn off); second and third antennal segments black ventrally (bb) |
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3 | First metasomal segment tricoloured ventrally (a); clypeus entirely yellow (b) or with small medial black patch; parategula yellow (c); [basal half of antenna usually extensively yellowish brown ventrally; pronotum broadly yellow posteriorly] |
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– | First segment bicoloured ventrally (aa); clypeus partly black (bb); parategula black (cc) |
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4 | Apical third of clypeus broadly yellow laterally (a); setae of mesoscutum comparatively short anteriorly (b); clypeus less convex compared to face in lateral view (c) |
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– | Apical third of clypeus broadly black laterally (aa); setae of mesoscutum comparatively long anteriorly (bb); clypeus more convex compared to face in lateral view (cc) |
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5 | Posterior part of first metasomal tergite comparatively slender in ventral view (a) and weakly convex dorsally in lateral view (b); apical lamella of second tergite often subhyaline or pale yellow (c), |
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– | Posterior part of first tergite comparatively robust in ventral view (aa) and more convex dorsally in lateral view (bb); apical lamella of second tergite dark brown or blackish (cc), rarely yellowish; setae of second tergite either long (dd) or inconspicuous in lateral view (ddd) |
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6 | Second metasomal tergite with three large (more or less separated) black spots medially (a); first metasomal segment largely orange or yellow (b); clypeus entirely yellow (c); mesoscutum with medium-sized to large yellow or orange patch laterally (d); [propleuron with short setae] |
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– | Second metasomal tergite with wide black band medially (aa); first segment mainly black (bb); clypeus at least medially partly black (cc); mesoscutum usually entirely black (dd) or with pair of small to large patches (large specimens of |
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7 | Second metasomal sternite with long setae (a), rarely intermediate; scape entirely black anteriorly (b), rarely with short yellow stripe; inner tooth of hind tarsal claw narrower and rather acute (c); clypeus comparatively deeply emarginate medio-apically (d); [hind tarsus dark brown; dorsal setae of scape often long; second tergite rather remotely punctate and bristly setose] |
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– | Second sternite with short to medium-sized setae (aa), rarely intermediate (Iberian Peninsula); anteriorly scape (except sometimes apical third or half) yellow (bb); inner tooth of hind tarsal claw wider and distinctly truncate (cc); clypeus less emarginate medio-apically (dd); but comparatively deep in |
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8 | Apical lamella of second tergite subhyaline or pale yellowish (a); ventral corners of clypeus narrower because of deeper medio-apical emargination (b); propodeum usually with medium-sized smooth interspaces (c) and anterior half of median groove distinct (d); second tergite comparatively convex medially and with short setae (e); first tergite slender in dorsal view (f); parategula frequently more or less yellow (g); labrum partly or entirely yellow (h); [hind basitarsus often brown basally] |
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– | Apical lamella of second tergite brownish or blackish (aa), rarely pale brown or pale yellowish; ventral corners of clypeus wider apically because of shallow medio-apical emargination (bb); propodeum usually without smooth interspaces (cc) and anterior half of median groove largely reduced (dd); second tergite less convex medially and with medium-sized to long setae (ee); first tergite less slender in dorsal view (ff); parategula usually black (gg) or largely so; labrum entirely dark brown or black (hh) |
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9 | Third tergite and sternite partly yellow posteriorly (a); hind basitarsus more or less darkened basally and outer apex of hind tibia with blackish patch dorsally (b); first metasomal tergite elongate in lateral view (c); first tergite usually less densely punctate subposteriorly (d); second tergite more shiny and usually less densely sculptured laterally (e); [large specimens (fore wing about 10 mm) have mesoscutum frequently with a pair of yellow patches antero-laterally; |
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– | Third tergite and sternite black posteriorly (aa); hind basitarsus and outer apex of hind tibia entirely yellow (bb); first tergite less elongate in lateral view (cc); first tergite more densely punctate subposteriorly (dd); second tergite more densely sculptured dorsally (ee) |
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10 | Middle of propodeum with small interspaces between coarse punctures micro-sculptured (a); mesoscutum with pair of yellow (and often large) patches antero-laterally (b); clypeus with coarser punctures, especially apically (c); length of fore wing 10–13 mm |
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– | Middle of propodeum either without interspaces between punctures (aa) or small interspaces present and smooth (aaa); mesoscutum entirely black or with smaller linear patches antero-laterally (bb); clypeus often only punctulate or with less coarse punctation, especially apically (cc); length of fore wing 7–11 mm |
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11 | Apical half of scape anteriorly (a) and clypeus (b) black or largely so; second sternite with narrow yellow band apically (c) |
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– | Apical half of scape anteriorly (aa) usually and clypeus (bb) partly yellow; second sternite with wider yellow band apically (cc) |
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12 | Pronotum flattened medio-posteriorly and narrowly yellow (a); apical corners of clypeus more protruding (b); third metasomal tergite entirely black (c); inner tooth of hind tarsal claws shorter and wider (d) and claw curved (e); [if second sternite with some long setae and first tergite comparatively slender, see |
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– | Pronotum convex medio-posteriorly and wider yellow (aa); apical corners of clypeus less protruding (bb); third tergite partly yellow (cc); inner tooth of hind tarsal claws slightly longer and narrower (dd) and claw less curved (ee); [if outer side of hind coxa mostly with short setae, shape of hind claws different, second tergite more punctate and fore tibia entirely yellow, compare with very similar |
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13 | Clypeus yellow medio-dorsally (a) and apical half largely black (b); posterior half of first tergite usually more robust in dorsal view (c); medio-posteriorly third sternite yellow (d); apical corners of clypeus slightly less protruding (e) |
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– | Clypeus black medio-dorsally (aa), if yellow (aaa) then apical half of clypeus also largely yellow (bb); posterior half of first tergite less robust in dorsal view (cc); medio-posteriorly third sternite usually black (dd) or yellow band reduced; apical corners of clypeus slightly more protruding (ee) |
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14 | Medio-posteriorly third metasomal sternite broadly black or dark brown (a); outer side of hind coxa mostly with very long setae in dorsal view (b); first tergite comparatively convex in lateral view (c); head conspicuously long setose (d); fore tibia often with dark brown or blackish patch medio-posteriorly (e); second tergite finely punctate (f); [yellow dorsal part of clypeus transverse |
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– | Medio-posteriorly third sternite yellow (aa) or narrowly interrupted; hind coxa with short to medium-sized setae in dorsal view (bb); first tergite less convex in lateral view (cc); head less conspicuously setose (dd); fore tibia entirely yellow (ee) or with blackish patch medio-posteriorly; second tergite more coarsely punctate (ff) |
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15 | Laterally propleuron regularly short setose in lateral view, setae curved and 0.1–0.3 times as long as occipital setae (a); hind tarsus (except dark brown telotarsus) brownish yellow (b); [apical lamellae of propodeum more or less darkened] |
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– | Laterally propleuron irregularly medium-sized setose in lateral view, setae straight (aa) or adpressed; hind tarsus dark brown (bb) or largely so |
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Males
16 | Setae of occiput short to medium-sized (a); second and third antennal segments reddish or brownish ventrally (b); [antennal hook distinctly bent; laterally pronotum often broadly yellow] |
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– | Setae of occiput long (aa; but sometimes worn off); second and third antennal segments black ventrally (bb) |
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17 | Antennal hook robust (a, b) and in lateral view claw-like (b); ventro-lateral corners of clypeus rather acute (c); [first tergite bicoloured ventrally; mesoscutal setosity short] |
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– | Antennal hook more slender (aa, bb) and less claw-like in lateral view (bb), but sometimes intermediate in SW Europe; ventro-lateral corners of clypeus slightly more obtuse (cc); [first tergite tri- or bicoloured ventrally] |
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18 | First metasomal segment tricoloured ventrally (a); antennal hook more slender in lateral view (b; sometimes less than shown); setae of eye incision at most half as long as apical width of scape in lateral view (c); parategula largely yellow (d), rarely dark brown; [antenna extensively yellowish brown ventrally as in |
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– | First segment bicoloured ventrally (aa); antennal hook less slender in lateral view (bb); setae of eye incision about as long as apical width of scape in lateral view (cc); parategula black (dd); [ |
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19 | Apical half of first metasomal segment largely orange or yellow dorsally (a); mesoscutum with medium-sized to large yellow or orange patch laterally (b); [propleuron with short setae] |
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– | Apical half of first segment largely black dorsally (aa); mesopleuron mainly or entirely black anteriorly (bb); mesoscutum black (cc) or with small transverse patch laterally (ccc), rarely larger |
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20 | Middle of propodeum with matt, micro-sculptured interspaces (a); mesoscutum with pair of transverse yellow patches antero-laterally (b), but sometimes reduced; apical antennal hook robust apically and dark brown (c); base of mandible often partly pale yellowish (d); length of fore wing 9–13 mm; [posterior half of first tergite very robust] |
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– | Middle of propodeum without micro-sculptured interspaces (aa); mesoscutum entirely black (bb); apical antennal hook slender apically and yellow (cc), rarely infuscate or dark brown ( |
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21 | Second metasomal sternite with some long setae (a), rarely medium-sized; posterior part of first tergite flattened in lateral view (b); scape black anteriorly or largely so (NW Europe; c), but more or less yellow in S Europe; ventro-apical corners of clypeus narrower (d); first tergite without pair of yellow spots (e) or spots minute; [second tergite conspicuously setose and remotely sculptured] |
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– | Second sternite with short or medium-sized setae only (aa); posterior part of first tergite convex in lateral view (bb); scape yellow anteriorly (c) or largely so; ventro-apical corners of clypeus wider (d); usually first tergite with pair of yellow spots (ee), but often reduced in |
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22 | Second tergite coarser punctate (a); first tergite slightly less robust in ventral view (b); third tergite more extensively yellow (c); setosity of scape either long dorsally (d) or (Balkan Peninsula) short |
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– | Second tergite comparatively finely punctate (aa); first tergite slightly more robust in ventral view (bb); third tergite less extensively yellow (cc); scape short setose dorsally (dd); [dark form with dark scape, more or less partly long setose second sternite and finely punctate second tergite] |
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23 | Antennal hook short, bent and slightly wider medially (a); apical lamella of second tergite subhyaline or pale yellow (b); second tergite more or less concave medio-posteriorly in lateral view (c); clypeus slightly more emarginated medio-apically (d) |
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– | Antennal hook longer, nearly straight (but medio-dorsally slightly depressed) and medially more slender (aa), but intermediate in |
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24 | Outer side of hind tibia with black or dark brown patch dorso-apically, contrasting with mainly pale hind basitarsus (a); first tergite comparatively slender in ventral (b) and dorsal (c) view; apical half of antennal hook with more or less dark brown keel dorsally and relatively slender (d); [second metasomal tergite shiny apico-laterally] |
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– | Outer side of hind tibia dorso-apically and hind basitarsus yellow (aa); first tergite usually more robust in ventral (bb) and dorsal (cc) view; apical half of antennal hook without dark brown keel dorsally (dd), if slightly developed then less slender; [second tergite usually with satin sheen apico-laterally] |
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25 | Third and fourth metasomal tergites black (a), rarely third tergite narrowly yellow; mesoscutum scrobiculate-reticulate (b); apical lamella of second tergite yellow or brownish (c); [antennal hook with submedial depression] |
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– | Third and fourth tergites partly yellow (aa); mesoscutum coarsely punctate or punctate-reticulate (bb); apical lamella of second tergite dark brown or blackish (cc) |
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26 | Setae of propleuron regular and 0.1–0.3 times as long as occipital setae (a); antennal hook nearly straight (a; sometimes slightly more curved than illustrated) |
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– | Setae of propleuron irregular and mainly straight and 0.3–0.7 times as long as occipital setae (aa); antennal hook more curved (bb) |
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27 | Antennal hook largely dark brown or brown (a), more slender and slightly bent (b); penultimate antennal segment dark brown ventrally (c); second metasomal tergite usually rather bristly setose (d) |
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– | Antennal hook yellow (aa) or largely so, less slender and more bent (bb); penultimate antennal segment yellowish brown ventrally (cc); second metasomal tergite usually only short setose (dd), but bristly in typical |
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28 | Outer side of hind coxa with short to medium-sized setae in dorsal view (a); first metasomal tergite usually slightly less convex in lateral view (b); head moderately setose (c); apical yellow band of second tergite less widened dorsally (d) |
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– | Outer side of hind coxa with very long setae in dorsal view (aa); first tergite comparatively convex in lateral view (bb); head conspicuously long setose (cc); apical yellow band of second tergite often comparatively wide dorsally (dd) |
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29 | Second metasomal tergite moderately to coarsely punctate (a) and usually rather dull (b); inner side of apical hook with small setae (c); [pale form of |
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– | Second metasomal tergite sparsely punctate or punctulate (aa) and shiny (bb); inner side of apical hook without distinct setae (cc) |
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The holotype female of
The dark typical form is mostly boreo-alpine of distribution and occurs in Scandinavia (up to S Finland and SE Sweden), UK (England and Wales) and mountainous regions in Central Europe (reaching 1550 m altitude in Switzerland). The pale typical form (= f.
The holotype female of
SE and C Europe, NW Asia. Examined specimens originating from Austria (type locality), Hungary, Czech Republic, and Slovakia (but most specimens from these countries in collections belong to
Males from the Balkan Peninsula have the apical half of antennal hook distinctly flattened, different from the wider apical half in Central European males (Fig.
Rather common in most of Europe (including southern Scandinavia;
North Africa, South Europe (*Spain, *Portugal, Italy (Sardinia, Sicily), *N Macedonia, *Bulgaria, *Greece) and *Turkey.
Rarely collected species in C and S Europe, but common in Spain and Portugal (
Asia, Central and South Europe. Absent in Switzerland according to
This species is in need of a critical revision; the few molecular data indicate that several cryptic species may be included under
Mediterranean, Balkan Peninsula, rarely in Central Europe (e.g., Switzerland only in Ticino and Valais and late in season (July–October;
Large specimens (fore wing length about 10 mm) have frequently a pair of yellow patches on the mesoscutum antero-laterally; the patches vary from minute to large. The photographs of the female holotype kindly supplied by Roberto Poggi (
Widespread in most of Europe, but considered absent from UK (only known as vagrant in England;
The depository of the female holotype of
Widely distributed in Europe but relatively rare in collections from NW and S Europe (e.g., only
The female lectotype of
One of the common species in S Europe, reaching Germany (but very rarely collected) and Belarus. Known from Corsica, Sardinia, Sicily, and Malta (
As shown by the short setae of the hind coxa, the robust posterior part of the first metasomal tergite and frequently present moon-shaped yellow patch of the clypeus in females of both
Examined specimens are from Spain, France, Bulgaria, and Turkey. This rarely collected species is also reported from Albania, Italy (type series), Greece and Ukraine (Crimea).
Similar to
An endemic species of Sardinia (Italy) and Corsica (France), occurring from sea level up to 1600 m altitude in Corsica (
Unfortunately, the original description does not include any remarks on the shape of the apical antennal segment of the male. The males should have the apical antennal segment narrower basally and less curved than in typical
A comparatively rarely collected species in C and S Europe as well in NW Asia. The typical form occurs in southern European Russia up to western Siberia. In Switzerland occurring between 255 and 1250 m altitude (
As pointed out by
Specimens in
C and S Europe, but unknown from Sardinia (
Conspicuous orange species only recently known to occur in Europe (
Central Asia, European Russia, Ukraine (Crimea).
We are very grateful for photographs of Fabricius, Kriechbaumer, Blüthgen and Guiglia types kindly supplied by Lars Vilhelmsen and Sree Gayathree Selvantharan (Natural History Museum of Denmark, Copenhagen), Stephan and Olga Schmidt (Zoologische Staatssammlung, München), Michael Greeff (Eidgenössische Technische Hochschule, Zürich) and Roberto Poggi (Museo Civico di Storia Naturale Giacomo Doria, Genoa), respectively.
We thank Frederique Bakker (